Classification, Phylogeny, and Zoogeography of the Genus Penlypus (Coleoptera: Cleridae)

GINTER EKIS

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 227 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION

Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world cf science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover.

S. Dillon Ripiey Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 227

Classification, Phylogeny, and Zoogeography of the Genus Perilypus (Coleoptera: Cleridae)

Ginter Ekis

MAR 3119771 i

SMITHSONIAN INSTITUTION PRESS City of Washington 1977 ABSTRACT Ekis, Ginter. Classification, Phylogeny, and Zoogeography of the Genus Perilypus (Coleoptera: Cleridae). Smithsonian Contributions to Zoology, number 227, 138 pages, 386 figures, 2 tables, 1977.—New World Genus Perilypus is redefined, and its natural history summarized. Species occur in at least three macrohabitats defined as oak, liana, and herbaceous assemblages. Beetles with rectangular bodies inhabit oak and liana assemblages whereas oval-bodied beetles are found among herbaceous plants; mimetic interactions are thought to be the basis for these habitat-body form correlations. Perilypus mimics are regarded as generalists having evolved in appearance and/or behavior characteristics that simulate more than one distasteful model. Altitudinally the genus ranges from sea level to 3400 meters; however, most specimens were collected between 1000 and 2000 meters. Laboratory observations indicate that Perilypus beetles are highly predatory, with size and rigidity of victims being important limiting factors in prey acceptability. The criteria used for recognition of species and of infra- and suprageneric groupings are discussed and techniques for dissections, illustrations, and measurements are described, a morphological analysis of the major external and internal organs is presented, and a key to the species groups and species based on adult characteristics is provided. Larval and pupal stages of Perilypus are described for the first time. The species groups are characterized, and given for each species are, as appropriate, synonymic list, diagnostic combination, description, discussion of structural and chromatic variation, discussion of natural history, geographic distribution, etymological derivation, locality records, general notes, and illustrations. Distribution maps with symbols visually convey the geographic range of species and species groups. There are 49 species presently recognized in Perilypus and 30 of these are described for the first time. Twelve new synonymies are established. A phylogeny of Perilypus is postulated using the Hennigian method of phylogenetic analysis. The zoogeography of the genus is discussed in terms of faunal limits and vicariance. Concepts of historical zoogeography ("faunal tracks", "center of origin and dispersal", and "forest refugia") are briefly discussed and two of them are used to explain distributions of extant Perilypus taxa. Relationship discussions within the species groups vary in completeness according to availability of data. A fairly complete treatise is presented for the larger limbatus, quadrilineatus, and ornaticollis groups, but the treatment of the reventazon group, the largest group in the genus, is very fragmentary. Ancestral Perilypus probably evolved in northern Central America, during mid-Tertiary, and ultimately evolved into eight species groups. Progenitors of the frontalis, criocerides, chaletoides, and quadrilineatus groups remained in northern Central America as did all of their descendants. Ancestors of the limbatus and viridipennis groups dispersed to southern Central America; some members of both groups ultimately penetrated South America. The ornaticollis group, the most derived species group in the genus, and the reventazon group are especially widespread geographically. [Abstract in Spanish on p. 135] OFFICIAL PUBLICATION DATF is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus). Library of Congress Cataloging in Publication Data Ekis, Ginter Classification, phylogeny, and zoogeography of the genus Perilypus (Coleoptera, Cleridae) (Smithsonian contributions to zoology ; no. 227) Bibliography: p. 1. Perilypus. 2. Insects—Classification. I. Title. II. Series: Smithsonian Institution. Smith- sonian contributions to zoology ; no. 227. QL1.S54 no. 227 [QL596.C62] 591'.08s [595.7'64] 76-18705 Contents

Page Introduction 1 Natural History S Macrohabi tats 4 Mimicry ; 4 Seasonal and Altitudinal Distribution 4 Laboratory Observations 4 Criteria for Species and for Infra- and Supraspecinc Groupings 5 Materials, Methods, and Terminology 6 Dissecting Techniques 6 Illustrations 6 Measurements 7 Types 7 List of Abbreviations 7 Morphological Analysis 8 External Morphology 8 Head 8 Prothorax 9 Mesothorax 12 Metathorax IS Wings IS Legs IS Abdomen 16 Reproductive Organs 18 Internal Morphology 18 Digestive System 18 Nervous System 20 Reproductive Systems 20 Genus Perilypus 20 Key to the Species of Perilypus 23 The frontalis Group 26 1. Perilypus frontalis (Gorhani), new combination 28 2. PerUypus sensilis, new species SO The criocerides Group SS 3. Perilypus criocerides (Gorham), new combination SS 4. Perilypus insectus, new species 35 The viridipennis Group 36 5. Perilypus viridipennis (Spinola), new combination 37 6. Perilypus relucens (Gorham), new combination 39 7. Perilypus levis, new species 42 The limbatus Group 45 8. Perilypus limbatus (Gorham), new combination 45 9. Perilypus columbicus (Spinoia), new combination 48 10. Perilypus apocopatus, new species 50 11. Perilypus iris, new species 52 12. Perilypus acus, new species 52 IS. Perilypus buga, new species 54 14. Perilypus latilira, new species 55 15. Perilypus prolixipenis, new species 58

iii IV SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Page 16. Perilypus testaceicornis (Pic), new combination 60 17. Perilypus decoris, new species 60 The reventazon Group 61 18. Perilypus reventazon, new species 63 19. Perilypus nigriventris (Gorham), new combination 64 20. Perilypus sapientis, new species 65 21. Perilypus claudus (Wolcott) 66 22. Perilypus cultratus, new species 69 23. Perilypus bilineatus (Gorham), new combination 71 24. Perilypus cerroazul, new species 73 25. Perilypus ordinatus, new species 73 26. Perilypus calcaris, new species 74 27. Perilypus orthopleuridus (Thomson), new combination 76 28. Perilypus exilis, new species 78 29. Perilypus latilobus, new species 79 30. Perilypus dislinctus (Chevrolat), new combination 80 31. Perilypus crassus, new species 82 32. Perilypus immitis, new species 83 33. Perilypus caliculus, new species 83 34. Perilypus carbonarius Spinola 86 The chaletoides Group 88 35. Perilypus chaletoides, new species 88 36. Perilypus bicolor (Chevrolat), new combination 91 The quadrilineatus Group 92 37. Perilypus quadrilineatus (Chevrolat), new combination 92 38. Perilypus telephoroides (Gorham), new combination 94 39. Perilypus coroniformis, new species 96 40. Perilypus fluctus, new species 98 41. Perilypus bicristatus, new species 99 The ornnticollis Group 99 42. Perilypus ventralis (Gorham), new combination 102 43. Perilypus ornaticollis (LeConte), new combination 105 44. Perilypus galbeus, new species 107 45. Perilypus prolixicornis, new species 108 46. Perilypus pilatus, new species Ill 47. Perilypus antarius, new species Ill 48. Perilypus sinuapicis, new species Ill 49. Perilypus floralis (Gorham), new combination 113 Immature Specimens 114 Phylogeny and Zoogeography 116 Phylogenedc Methods 116 Zoogeographic Methods 125 Phylogenetics and Zoogeography of Perilypus 126 Intergeneric Relationships 126 Relationships among, the Species Groups 126 Relationships within the Species Groups ISO Abstract in Spanish 135 literature Cited 136 Classification, Phylogeny, and Zoogeography of the Genus Perilypus (Coleoptera: Gleridae)

Ginter Ekis

Introduction the beginning stages of this work; to resolve the species concept in Perilypus, clarify nomenclatural Checkered beetles of the genus Perilypus are ex- problems, and provide diagnostic aids for species clusively New World and predominantly tropical; recognition. At first there was little more to be done they are predaceous surface dwellers with great with this group because only a handful of speci- agility in gait and flight. Structurally, the beetles mens were available for study. Perilypus specimens are either rectangular or oval in body form and are not abundantly collected in nature, as is the their integumental color ranges from dull black to floricolorous patterns of yellow, red, and blue. case with most mimics; consequently, the beetles Woody vegetation in dry montane forests seems to were unfamiliar to most keepers of collections and be the macrohabitat for many of the species, but essentially unavailable for loan shipments. In time, lampyroid mimics occur on herbaceous plants lo- additional specimens became available, particularly cated in the more open highland habitats. Intra- during my tenure as a research fellow of the Smith- and interpopulation color polymorphism (often sonian Institution. During that period (June 1973 correlated with sex) and mimetic life styles are some to June 1974) two research grants provided financial interesting traits of most of the species. support for an extensive field expedition to Central I first acquired an interest for these beetles in and South America and a visit to various museums 1970 when William F. Barr of the University of in Europe. These expeditions nearly doubled the Idaho conveyed the following notes about speci- available study material. Particularly important was mens of Perilypus (heretofore in the genus Coly- the trip to Latin America, during which I obtained phus Spinola) sent to him for identification: "I'm data about the natural history of the group, col- at a loss as to what to call these Colyphus speci- lected specimens of several species in fluid for mens. The names available in the literature really analyses of internal anatomy, and discovered the are meaningless until we can establish what con- larval and pupal stages of the genus. stitutes a species in this group. The possibility of a The procurement of additional specimens and considerable amount of variation within a species biological data prompted an expansion of the scope including sexual dimorphism should not be over- of this study to include generalizations about looked." From this reply evolved my objectives for Perilypus natural history, and analyses of internal and external anatomy, and of phylogeny and zoo- Ginter Ekis, Research Fellow, Department of Entomology, geography. The reader may find some of these National Museum of Natural History, Smithsonian Institu- sections less than complete; for example, in the section, Washington, D.C. 20560. tions of phylogeny and zoogeography, relationships SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 1-2.—Perilypus decoris: 1, forebody: 2. habitus. NUMBER 227 within the largest species group, the reventazon theory. John M. Kingsolver kindly made avail- group, remain undeciphered. Even a terse and/or able a microscope during the initial stages of provisional treatment of some of the aforemen- this work. I also must acknowledge his patience tioned subjects, however, may positively influence a during repeated visits to his office, the first point group's systematic progress. The purpose of this of inquiry whenever I needed equipment relevant paper is twofold. First, it is offered as a unit of to taxonomic work. As usual, William F. Barr of information consisting of all the current knowledge the University of Idaho was very helpful in making about the genus Perilypus, including a resynthesis available his notes and photographic transparencies of the data in sections of morphology, practical of type specimens, and George C. Steyskal rendered taxonomy, and evolutionary relationships. Second, guidance on matters relevant to nomenclature. I am this work is intended to establish a ground plan for indebted to George L. Venable for drawing the revisions involving genera of the subfamily Clerinae. habitus and forebody illustrations of P. decoris, LITERATURE REVIEW.—Perilypus was validated by new species (Figures 1, 2) and to Michael W. Spinola in 1841; he included the name in a diag- Druckenbrod for illustrating P. limbatus larva and nostic key of the clerid genera of the world. The pupa (Figures 370, 380,381). genus was subsequently more fully described by I extend my gratitude to the museum curators Spinola in 1844a on the basis of P. carbonarius and private collectors (see "Materials") who en- which until recently was classified under the sub- trusted me with their specimens. For lending type family Tillinae. In a previous paper, I (1975:25) specimens I am grateful to Christine M. F. von discussed the classification of the genus and trans- Hayek (British Museum of Natural History), Mme. ferred two other Spinola species to Perilypus from A. Bons (Museum d'Histoire Naturelle de Paris), Colyphus. In this work, the majority of Colyphus F. Heike (Museum fur Naturkunde on der Hum- species described by other authors are also trans- bolt-Universitat zu Berlin), Marco Spinola (Tas- ferred to Perilypus. There is little else to say about sarolo Castle in Novi Ligure, Italy), Henry Dybas the history of the group, except that the species (Field Museum of Natural History), and John F. were never studied collectively, and that species Lawrence (Museum of Comparative Zoology). descriptions are scattered throughout the literature, This study was supported in part by a one year are very brief, and rarely include illustrations. Be- Smithsonian research fellowship in the Institution's sides Spinola, the following authors published one program in Evolutionary and Systematic Biology, a or more names of species now considered valid and travel grant from the American Philosophical So- currently placed in Perilypus: Chevrolat (1843, ciety (Penrose Fund #6658) for type studies abroad, 1874), Gorham (1878, 1882, 1886), LeConte (1880), and a Walter Rathbone Bacon Scholarship from the Pic (1941), Thomson (1860), and Wolcott (1927a). Smithsonian Institution (awarded to T. L. Erwin) ACKNOWLEDGMENTS.—During the course of this for field studies in Latin America. For typing vari- study, I was very fortunate to work at the National ous sections of the manuscript I am grateful to Jo Museum of Natural History, Smithsonian Institu- Whitehead and Nancy Stefansson. Jos6 Cuatrecasas tion, with many competent persons who were al- kindly translated the abstract into Spanish. ways willing to discuss concepts or assist in material needs. For practical reasons, only a few can be men- Natural History tioned herein. I am particularly indebted to Terry L. Erwin, who provided among other things, mu- Although little is known about the natural his- seum space, equipment, travel funds for fieldwork tory of Perilypus beetles, some general comments in Latin America, a review of the manuscript, and, can be made about their macrohabitats, approxi- most important, guidance and encouragement dur- mate seasonal and altitudinal distribution, and ing a research fellowship at the Smithsonian In- apparent involvement in several mimetic com- stitution. This manuscript was also reviewed by plexes. More specific notes relevant to mating be- Donald R. Whitehead of the University of Michi- havior and prey acceptance are based on laboratory gan and Lee H. Herman, Jr., of the American observations. In addition, on page 47 I briefly Museum of Natural History, with whom I also allude to the larval and pupal microhabitat of P. enjoyed occasional discussions about systematics limbatus. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MACROHABITATS bers may simulate lampyrids, cantharids, lycids, or chrysomelids. Characteristics of the mimic impor- Field observations (June, July) in Mexico, Guate- tant in the perfection of the mimetic pattern may mala, Costa Rica, and Venezuela indicate that spe- involve body form, integumental color, and be- cies of Perilypus frequent at least three kinds of havior, or a combination of all three as in P. vegetation assemblages, here broadly denned as the ventralis. The mimetic relationships within Perily- oak assemblage, the liana assemblage, and the her- pus will be discussed in more detail. For the present baceous assemblage. Specimens were collected from suffice to say that there is a tendency among some the oak assemblage, essentially a montane oak-pine species of Perilypus to simulate more than one forest, by beating low hanging branches of Quercus. distasteful model; that is, the mimic has evolved a The liana assemblage consists of an assortment generalized appearance and/or behavior that simu- of lianas intertwined with branches, twigs, and low lates members of several model species with more or canopy shrubby vegetation. A substantial part of less similar color patterns. Membership in such a the woody vegetation is dead. The most productive generalized mimetic complex requires phenotypic method of collecting specimens from the liana as- plasticity on the part of the mimic. Perhaps the semblage is by beating between 10:00 AM and 1:00 extensive color polymorphism present in this genus PM. This macrohabitat, usually found in the drier is a manifestation of that requirement. regions of montane forests (about 1500 m), also is frequented by wood-boring beetles. This suggests, at least superficially, a potential biological relation- SEASONAL AND ALTITUDINAL DISTRIBUTION ship between the wood borers and the species of Seasonal and altitudinal distribution records were Perilypus found among them; under laboratory taken from collector labels and from data recorded conditions, bark bettles were the preferred food of by me during field work. Most adult specimens were P. limbatus. collected from June to August; very few were ob- The herbaceous assemblage involves broad-leaf tained before June or after August, and none dur- herbaceous plants (including such members of Com- ing December or January. Three immatures (two positae as Vernonia deppeana Lessing), which were larvae and one pupa) were captured in June. The particularly evident along hedgerows adjacent to composite altitudinal range of the members of cultivated fields and near roadsides. Adult Perilypus Perilypus extends from sea level to about 3400 on these plants were collected easily by hand dur- meters, most specimens being collected between ing late afternoon hours. 1000 to 2000 meters. There is an interesting correlation between the body shape of the beetles and two of the vegetation types on which they were collected. Beetles with LABORATORY OBSERVATIONS rectangular bodies were found in liana assemblages, Specimens of P. limbatus were collected alive, whereas those with oval bodies were found on sexed, and maintained in petri dishes for general broadleaf herbaceous plants. Consequently, unlike observation. Longevity of captive specimens ranged the slender beetles from the liana plants, the more from five to ten days after capture; males perished ovate beetles always were exposed and conspicu- first and became a meal for survivors if not quickly ously visible on broad-surface foliage. This is not removed from the enclosures (dried specimens were surprising, however, as oval Perilypus are lampyrid not eaten). mimics whose success is dependent on their "incon- Interphenon matings occurred between members spicuous" exposure among models. of all phena (pp. 5, 47). The mating position is dorsoventral. The male lifts his forebody onto the MIMICRY female while maintaining his hind legs in contact with the substrate; copulation lasts approximately Even a cursory survey of Neotropical clerids re- 45 seconds. Precopulation events are rapid and un- veals how extensively mimicry and warning colora- complicated. The male overtakes the female from tion have influenced clerid evolution. These traits behind and with his hind legs on the substrate certainly are conspicuous in Perilypus whose mem- clasps the female epipleural margin with his pro- NUMBER 227 and mesotarsal claws. I suspect that the convex characteristics are exceptionally homogeneous at female epipleuron and the conspicuously asym- the infraspecific level but otherwise they are highly metrical male protarsal claws function as copulatory variable. Characteristics of the ovipositor are spe- anchorage structures. cifically distinct within some species groups. Field and laboratory observations suggest that I consider stable gaps (morphological or other- only one egg is released during each oviposition. wise) not correlated with sex between sympatric The habit of depositing a single egg at each ovi- forms to indicate nonconspecificity; I assume such position site is possibly a safeguard against larval gaps are manifestations of disrupted gene flow. cannibalism. One captive female devoured her Allopatric samples are regarded as conspecific if singly laid egg immediately after it was deposited they show no appreciable morphological or biologi- on the surface of a small twig. cal differences. Within this definition, minor varia- My observations pertinent to prey acceptability, tions are interpreted as due to geographic variation. admittedly based on superficial tests, show that When allopatric samples are notably different, how- specimens of P. limbatus feed on a wide array of ever, the magnitude of their difference is compared insects as most checkered beetles probably do. Ap- with that observed between sympatric species. In parently two important limiting factors in prey absence of contrary evidence and when such differ- acceptance are size and rigidity of the exoskeleton ences between the allopatric samples approximate of the potential victim. Smaller prey, such as those between the sympatric species, I judged allo- scolytid beetles, are held with the middle legs (usu- patric forms not conspecific. ally around the abdomen) and simultaneously I generally oppose formal naming of species sub- oriented with the front legs. The predator follows units, particularly in the absence of nonanatomical a distinct sequence when devouring its catch: first information. Even when such information is avail- eaten are the contents of the cervical membranes, able, it usually consists of gross approximations of followed by the soft tissues at the promesothoracic environmental limits and of distribution, and there- junction, and finally the contents of the abdominal fore is deficient for in-depth assessments of popu- cavity. Some larger beetles such as elaterids, chry- lation dynamics. The recent work of Ehrlich, et al. somelids, and cerambycids also were eaten, but (1975) and of authors cited therein demonstrates these were acceptable only when freshly killed and the level of experimentation necessary to gain with some exposure of internal tissue. meaningful information about interpopulation gene flow. Criteria for Species and for Infra- In addition to avoiding the introduction of am- and Supraspecific Groupings biguous trivial names into the literature, nonusage of the subspecific category necessitates a more thor- I accept the biological species concept (Mayr, ough discussion of interpopulation differences. Too 1969:26) as the theoretical premise for recognition often, and after minimal assessments, taxonomic of species despite the fact that conjectures about entities are named with justifications for their for- reproductive isolation are usually based on indirect mal recognition ill-defined or hidden under the evidence (in part indicative of our inability to de- cloak of theory. termine extent of gene flow). No species concept I believe that most differences among infraspecific alternative to Mayr's is more consistent with evolu- populations can be conveyed effectively and empha- tionary theory. sized through discussions, and if necessary, variant The nominal species included in this work repre- populations may be identified by informal desig- sent my best estimates (hypotheses) of what the bio- nations such as "morph" and "phenon." In this logical species of Perilypus are. Each estimate is work, noteworthy infraspecific variations are dis- based largely on anatomical criteria, but structural, cussed under "Variation" in the species descriptions. distributional, and biological characteristics are When variation is noted at the interpopulation integrated in species delimitations whenever pos- level, specimens exhibiting the atypical character- sible. In Perilypus the genital organs, and particu- istics are assigned under a "morph" name (e.g., red- larly the aedeagus, provide the most consistent legged morph). I assigned groups of specimens characteristics for species recognition. Aedeagal exhibiting a common intrapopulational color varia- SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY tion to a "phenon"; observed color polymorphism is may be dry mounted and subsequently rehydrated often sex dependent. to an extent that delicate internal tissue can be I do not formally recognize supraspecific taxa easily dissected. The quality of these dissections was within Perilypus. Until relationships among the often equivalent to those made with specimens various New World clerine groups (of dubious retained in fluid. generic rank) closely related to Perilypus are better Fluid-preserved specimens were dissected in water understood, I will recognize the "species group" as follows: First, the hind pair of wings was re- category and, when necessary, the "species sub- moved with microscissors, submerged in absolute group" category. Informal species groups and spe- ethanol, unfolded between two glass slides, dried, cies subgroups will identify adequately the respec- removed from the slide, mounted on paper point, tive major and minor monophyletic lineages of these and pinned beneath the specimen and at right genera until such time when a balanced generic angles to it. The second step involved removal of classification in the Clerinae is established. Lind- the abdominal contents including the copulatory roth (1969:xxiv) and Whitehead (1972:140) also organs. With an eye knife the abdomen was incised have alluded to the pitfalls of indiscriminate use of from the third spiracle to the anus; with a watch- the subgenus category. maker's forceps the digesting tract was severed at the base of the abdomen; and the intact mass of Materials, Methods, and Terminology internal organs was lifted out of the abdominal cavity. The organs were subsequently separated and I examined more than 1200 adult specimens, one reintroduced into Pampel's fluid until they could pupa, and two larvae of North, Central, and South be studied and illustrated. Once dissected, beetles American Perilypus, and approximately 300 adults were rinsed in water, dried on tissue paper, and and 12 larvae of other genera of Clerinae. The ma- pointed. Whenever possible, at least four fluid- jority of these specimens were loaned to me by preserved specimens (two males and two females) museums but material from private collectors is were dissected as described above, but the genital also well represented. I collected these insects from organs were extracted from all male specimens ir- Mexico southward to Venezuela and Charles W. respective of preservation technique used. O'Brien, Terry L. Erwin, and Henry P. Stockwell To remove membranous wings from a dry speci- made special efforts to collect material in Pampel's men, it was boiled for five minutes in water con- fluid especially for this study. taining five drops of 409 detergent. After dissection the beetle was washed in an ultrasonic cleaner, DISSECTING TECHNIQUES dried, and pinned. Extraction of genital organs from dry-preserved beetles involved severing the Fluid-preserved specimens were killed and fixed abdomen from the specimen, boiling in 10 percent in Pampel's fluid (glacial acetic acid, 4 parts; 40% KOH, and dissection procedures as described in the formaldehyde, 6 parts; 95% alcohol, 15 parts; water, previous paragraph. Occasionally internal repro- 30 parts). I found this preservative superior to any ductive organs from dry-preserved beetles were other that I have used to date (Ekis and Gupta, sufficiently rehydrated so that a detail study of 1971:52); the beetles are not discolored, do not be- them was possible. Genitalia and portions of in- come brittle internally, and can remain in fluid ternal organs were stored in glycerine and intro- for several years without notable damage. Several duced into plastic genitalia vials, which were structures described under "Morphological Analy- sealed with polyethylene stoppers and pinned below sis" would not have been noticed had I only studied appropriate specimens. dried-preserved specimens. Indeed, I found that by examining fluid-preserved specimens in fluid (usually water) integumental regions, and particularly ILLUSTRATIONS the soft membranous structures, were more clearly All drawings were prepared with a camera visible and easier to maneuver for detailed study. lucida mounted on a Wild stereoscopic microscope. Another advantage to collecting specimens in Pam- Sometimes line drawings of small organs (e.g., pel's fluid is that specimens collected in that fluid mouthparts) had to be optically magnified so that NUMBER 227 structural details, examined through a compound Colyphus melanopterus Dury. Because male genital microscope, could be presented in adequate scale. characteristics provide the most reliable criteria for In the drawings of the male internal reproductive diagnosing species, whenever possible I selected organs only one-half of the accessory glands, testes male specimens to establish holotypes or designate and associated tubes were drawn. The posterior lectotypes. Procedures pertinent to selection of pri- accessory glands are stippled for contrast. Unless mary or secondary types were carried out in accord- indicated otherwise the scale line accompanying a ance with the International Rules of Zoological drawing equals 1 mm. The scales for electron Nomenclature (1964). micrograph figures are presented in the legends. Data relevant to type-specimens and to type- Distribution maps, usually by species groups, are localities are noted at the beginning of each species provided; dots and various other symbols represent description. locality records. I used a question-mark symbol whenever the locality records referred to a country LIST OF ABBREVIATIONS only. All illustrated morphological structures discussed in the text are indicated by italics at first The following abbreviations are used in the text. mention. They indicate the repository of specimens and collections from which material was borrowed. Ad- MEASUREMENTS dresses and the names of curators within the institution, and private collectors are supplied to The meristic data provide a close approximation facilitate inquiries regarding the collections. of the size and proportions of the beetles and of AMNH American Museum of Natural History, New York, some of their parts. They also provide a quantita- New York 10024; L. H. Herman, Jr. tive basis for substantiating descriptive terms, such BMNH British Museum (Natural History), Entomology, as "feebly transverse," "boldly transverse." When SW7 5BD, London, England; Christine M. F. von available, I measured at least 10 males and 10 Hayek females of each species. All measurements were CASC California Academy of Sciences, San Francisco, made with an ocular micrometer through a Spencer California 94118; D. H. Kavanaugh CBAZ C. Bordon Azzali, Avenida Lima, Edificio Taor- stereoscopic microscope. Measurements and their mina, Apt. 8, Los Caobos, Venezuela; private abbreviations (when relevant) found in the descrip- collection tions are: length = body length, sum of lengths of CISC California Insect Survey, Entomology, University head, pronotum, and elytron (head length is the of California, Berkeley, California 94720; J. A. linear distance from the antennal carina to the Chemsak most posterolateral limit of the eye; lengths of CMPP Carnegie Museum, Insects and Spiders, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15212; pronotum and elytron are explained below); G. E. Wallace width = body width, maximum width across CMNH Cincinnati Museum of Natural History, 1720 elytra; AL = antennal length, sum of lengths of all Gilbert Ave., Cincinnati, Ohio 45202; C. Oehler articles; EL = elytral length, linear distance from CNCC Canadian National Collection of Insects, Entomol- the humeral margin, proximal to the suture, to the ogy Research Institute, Ottawa, Ontario, Canada; D. Bright, E. C. Becker elytral apex; EW = maximum width across the CUNY Cornell University, Entomology, Ithaca, New York elytron; HW = maximum width of the head across 14850; L. L. Pechuman eyes; IOW = maximum width between ocular su- DEIE Deutsches Entomologisches Institute, 13 Ebers- tures; PL = maximum dorsal pronotal length; walde, D. D. R.; H. Morge PLS = maximum lateral pronotal length; PW = FMNH Field Museum of Natural History, Chicago, Illi- maximum dorsal pronotal width. nois 60605; H. Dybas GEKI G. Ekis, c/o National Museum of Natural History, Smithsonian Institution, Entomology, Washington, D.C. 20560; private collection TYPES GHNE G. H. Nelson, Kansas City College of Osteopathy and Surgery, Anatomy, 2105 Independence Avenue, I examined the primary type-specimen (holotype Kansas City, Missouri 64124; private collection or lectotype) of each Perilypus species except HFHO H. F. Howden, Carleton University, Biology, Derestenus columbicus var. reynoldsi Wolcott and Ottawa, Ontario, Canada; private collection 8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

HPST H. P. Stockwell, Gorgas Memorial Hospital, Canal comprehensive morphological analyses in revision- Zone; private collection ary works. By investigating more than the con- ISNB Institute Royal des Sciences Naturelles de Bel- venient integumental and internal characteristics, gique, Bruxelles the basis for evolutionary inferences is maximized. LACM Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California With this in mind I decided to present a reasonably 90007; J. P. Donahue thorough morphological analysis of Perilypus and LMDR L. M. Drukenbrod, 5519 Decatur St., Bladens- to accomplish this I chose P. reventazon since its burg, Maryland 20710; private collection members exhibit the "typical" exoskeletal char- MCZC Museum of Comparative Zoology, Harvard Uni- acteristics of most Perilypus species. Herein, I treat versity, Cambridge, Massachusetts 02138; J. F. Lawrence most of these characteristics and those of the ali- MHNM Museu de Historia Natural de la ciudad de mentary canal, ventral nerve cord, and internal Mexico, Apartado 18-845, Mexico, D.F. 18; D.F.A. reproductive organs of both sexes. The characters Barrera and their states are defined and illustrated, not only MNHB Museum fur Naturkunde on der Humbolt- Universitat zu Berlin, 104 Berlin, Invalidenstrasse to facilitate the descriptions and discussion included 43, DDR; F. Hieke in this work, but also to provide a foundation of MNHP Museum National d'histore Naturelle, Entomol- terminology for future works. ogie, 45 bis, Rue de Button, Paris (Ve), France; A Bons, M. Villiers, J. J. Menier MSPI Massimiliano Spinola Collection, Tassarolo Castle, EXTERNAL MORPHOLOGY Novi Ligure, Italy; M. Spinola MZSP Museu de Zoologia da Universidade de Sao Paulo, Head Caixa Postal 7172, 04263, Sao Paulo, Brazil; H. Reichardt CRANIUM.—Snodgrass (1935:111) used head su- MZSI Museo ed Instituto di Zoologia Sistematica della tures to delimit six major areas of the generalized Universita di Torino, 10123 Torino, Via Giovanni insect cranium. Unfortunately, comparable areas Giolitti, 34 Italy; M. Zunino NMNH National Museum of National History, Smith- cannot always be so rigidly denned; in beetle adults sonian Institution, Entomology, Washington, D.C. some of the sutures described by Snodgrass are 20560; R. E. White, P. J. Spangler obscured or absent. An alternative used herein is RNHN Rijksmuseum van Natuurlijke Historie, Raam- to subdivide the cranial surface on a more general steeg 2, Leiden, Netherlands; J. Krikken basis. SEMK Snow Entomological Museum, University of Kan- sas, Lawrence, Kansas 66044; G. W. Byers The major structures and areas on the front of TAMT Texas A & M University, Entomology, College the head are illustrated in Figure 3. The fronto- Station, Texas 77843; H. R. Burke clypeal region is bounded laterally by the antennal TTUT Texas Tech University, Entomology, P. O. Box carina and interocular portion of the ocular suture, 4169, Lubbock, Texas 79409; D. Foster anteriorly by the labral base, and posteriorly by the UCMV Universidad Central de Venezuela, Instituto de Zoologia, Facultad de Agronomia, Agricola, Mara- hind limit of the interocular depressions. The epi- cay, Venezuela; G. Yepes T. stomal suture traverses the frontodypeal area, divid- UZMD Universiteits Zoologiske Museum, Universitet- ing it into the clypeus and the broader frons. The sparken 15, Copenhagen, Denmark; Sv. G. Larsson latter is marked by a pair of interocular depressions WBIV W. Bivin, Box 1854 Balboa, Canal Zone; private that partially encircle the frontal umbo. The region collection from the posterior limit of the frons to the post- WFBA W. F. Barr, University of Idaho, Entomology, Moscow, Idaho 83843; private collection occipital suture (Figure 7) is the epicranium (Figure ZMAN Zoologisch Museum der Universiteit van Amster- 3) which extends from the most dorsal aspect of the dam, Entomologie, Plantage Middenlaan 64, Am- ocular suture to the base of the epicranial acumina- sterdam 1004, Netherlands; J. P. Duffels tion (Figure 4). The gena (Figure 5) is the postocular region of Morphological Analysis the cranial wall, bounded posteriorly by the postoccipital suture, dorsally by the epicranium, and In studies of insect systematics, anatomy often ventrally by the gular suture and obscured lateral provides most if not all of the data accessible for borders of the submentum. The gena is conspicu- relationship discussions, hence the importance of ously sculptured with shallow wrinkles at the sides NUMBER 211 9 proximal to the ocular suture. The transverse gula and prominently visible in repose. Its outer facies is (Figure 5) is prolonged posteriorly into an oblong, broadly depressed and setose in the basal two-thirds. feebly elongate gular process that is microsetiferous Three dentes occupy most of the masticatory sur- on the hind margin. The free borders of the process face; the anterior dens is broadly arcuate and unite with the ventral portion of the inflexed acuminate; the medial dens is short, truncate, and postocciput. The latter borders the occipital fora- feebly arcuate; the posterior dens is broad, shallow, men and is widely splayed proximal to the epi- and projects an asperate profile. Behind the cranial acumination. The compound eyes are setose, posterior dens is a small transparent lobe, which boldly convex, have a well-developed ocular notch appears continuous with the mandibular penicillus; (Figure 6) in front, and consist of ommatidia that the latter extends anteriorly along the inner base are about as wide as the ocular suture. of the medial and posterior dentes. The condyle ANTENNA.—The antenna (Figure 8) consists of 11 and fossa of the mandibular base articulate with the articles, is serrate along the anterior border, and ventrolateral edge of the cranium. via the articulatory process of the scape pivots on The maxilla (Figure 12) connects to the cranium the antennifer (Figure 6) of the antennal sclerite. behind the mandibular fossa. Its triangular base, The scape is feebly arcuate and slightly concave the cardo, is bifurcated into a posterior inner and ventrally, and is about twice as long as the sub- outer cardal process. Anteriorly the cardo connects spherical pedicel. The articles of the flagellum are to the basistipes, a triangular sclerite flanked by the compressed dorsoventrally but as a group become palpifer and mediostipes. The latter is continuous progressively more subcylindrical toward the anten- with the lacinia, whose base is distinguishable in nal apex. The distal margins of articles 1 through maxillary dorsal view only. Distally the lacinia is 10 are truncate whereas article 11 is acuminate; all divided into a triangular medial lobe (the medio- articles are densely setose, particularly the distal lacinia) and smaller basolateral lobe (the five. The macrosetae (Figure 19) are long, acumin- laterolacinia). A short, transverse inflection divides ate, and grooved longitudinally, whereas the micro- the galea into a small basigalea and vastly larger setae are short, smooth, and blunt apically. distigalea. The maxillary palpus consists of four Microsetae occur only on the last five articles, be- setose articles; the first (the basal) is the shortest, ing concentrated on the dorsodistal region of ar- the second and third together are slightly longer ticles 7 to 10, and are particularly prominent on the than the fourth, and the fourth is cylindrical and dorsal surface of article 11. The ventral surface of membranous apically. the antenna is more sparsely setaceous, particularly The labium (Figure 13) connects to the submental at the distal third where microturbercles are pres- region of the cranium (Figure 5). Its mentum is ent. These tubercles are most abundant on the predominantly membranous, feebly sclerotized venter of article 11, and are very similar to the basally and laterally. The palpigers also are feebly sensilla basiconica of Tenebrio molitor Linnaeus sclerotized, subcylindrical, and partially underlie reported by Doyen (1966:110). the mentum. The ligula has a setose submarginal MOUTHPARTS.—The labrum (Figure 9) is a trans- fringe anteriorly, some scattered setae medially, verse, strongly emarginated preoral flap attached to and one exceptionally long setae proximal to each the frontoclypeal inflection. The labrum is vested bisetose palpiger. The labial palpus consists of three dorsally with long setae and has an anterior fringe articles; the first is conical; the second slightly of short setae. The torma projects from the hind swollen apically and nearly twice as long as the first; angles of the labrum and bifurcates distally into a and the third, the largest, is securiform, membra- medial tormal process and a lateral tormal process. nous along its distal border, and densely vested with The tormal sclerite is similarly bifurcated in other short recumbent setae. clerids (Solervicens, 1973a: 165; 1973b:235, 238) and in other beetles (Evans, 1961:307; Doyen, 1966:111). Prothorax Anteroventrally, the labrum has a U-shaped ridge that partially outlines a thin, feebly concave sdero- The prothorax (Figures 14-16) is a squat moder- tized region. ately transverse cylinder flexibly connected to the The mandible (Figures 10, 11) is subfalcifonn cranium by the cervical membrane. The cervical 10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

EPICRANIUM GENA COMPOUND IYES OCULAR SUTURE FRONS FRONTAL UMIO ANTENNAL INTEROCULAR CARINA DEPRESSION CLYPIUS EPISTOMAL SUTURE

SUBMENTAL REGION GULA GULAR SUTURE GULAR PROCESS GENA

EPICRANIAL ACUMINATION

OCULAR NOTCH

POSTOCCIPITAL SUTURE OCCIPITAL FORAMEN

POSTOCCIPUT

ANTENNIFER

FIGURES Z-8.~Perilypus reventazon: 3, head, anterior; 4, head, dorsal; 5, head, ventral; 6, head, lateral; 7, head, posterior; 8, antenna. (Soles = 1 mm) NUMBER %Z 11

MEDIAL TORMAL PROCESS LATERAL TORMAL PROCESS

^LATEROLACINIA J LAONIA

LABIAL PALPUS

PRONOTAL ARCH

UTUM ANTERIOR FORAMINAL FLANGE PROSTERNUM

CRYPTOSTERNUM PUURAl APOPHYSIS TROCHANTIN INTERCOXAL PROCESS ANTEROVENTRAL MEMBRANEOUS TUBERCLE POSTEROVENTRAL MEMBRANEOUS TUBERCLE POSTERIOR FORAMINAL FLANGE PROSTERNAL APOPHYSIS

FIGURES 9-18.—Perilypus reventazon: 9, labium; 10-11, mandible; 12, maxilla; 13, labium; 14, prothorax, dorsal view; 15, prothorax, lateral view; 16, prothorax, ventral view; 17, mesoscutellum, dorsal view; 18, mesoscutellum, lateral view. (Scale = 1 mm) 12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY sclerites are slender, feebly pigmented plates that dominantly glabrous; setae are present on the do not articulate with the head or prothorax. The anterior margin and on the feebly expanded distal dorsal and lateral wall of the prothorax comprise region of the intercoxal prosternal process. the pronotum; the ventral wall, or prosternum, is greatly reduced externally but substantially Mesothorax inflected. The pronotum (Figures 14, 15) is traversed by The pro-mesothoracic intersegmental membrane anterior and posterior depressions that delimit contains one dorsal and two ventral pairs of three regions, the pronotal arch, the pronotum transverse membranous tubercles (Figure 16). The proper, and the pronotal collar. At about the dorsal pair is similar in form and vestiture to the anterior fourth, the subapical depression marks the anteroventral pair. The posteroventral tubercles hind limit of the convex pronotal arch that extends differ from the others by being more conical and over most of the epicranial region when the head by bearing stout vertical setae; they are probably is reposed. To minimize the potential friction be- sensory organs as suggested by their connection tween the epicranial surface and the inner wall of with a mesothoracic nerve. the pronotal arch, the latter is lined with a trans- The mesothorax (Figures 21, 22) is approximately parent inflection, the anterior foraminal flange one-half the size of the metathorax. Its thick, (Figure 16). This thin, almost membranous flange strongly sclerotized notal plate (Figures 17, 18) may extends posteriorly to the inner ridge of the sub- be divided conveniently into scutum and scutellum; apical depression where it connects to the cervical the former is transverse, the latter subquadrate. A membrane. longitudinal mesonotal suture divides equally the In dorsal view the pronotum proper is subspher- rugose scutum, which partially projects into the ical, boldly convex laterally, sinuous anteriorly, pronotum. Side margins of the mesonotum are and truncate posteriorly. Near each hind angle broadly inflexed, particularly so below the scutel- there is a spheroid muscle-attachment depression, lum where the inflection serves as a shelf on which the pronotal fovea (Figure 14). Behind this fovea, rests the mesobasal angle of the elytron. the prebasal depression precedes the narrow pro- The mesopleuron (Figures 21, 22) is divided into notal collar, which widens at the sides and is a large triangular mesepisternum and a smaller, broadly inflected along its posterior margin. This tapering mesepimeron. The first is coarsely rugose inflection, the posterior foraminal flange (Figure in the dorsal half, conspicuously ridged at the 16), is a broad, rigid lining that combines with anterior margin, and is copiously vested with recum- the pronotal collar to provide additional rigidity to bent setae, as are most of the metathoracic pleural the posterior region of prothorax. Anteriorly, the and sternal plates. The substantially narrower flange connects to the ensuing intersegmental membrane. mesepimeron is finely rugose and emarginated at the pleurocoxal articulation. The prosternum (Figure 16) is a narrow trans- The mesosternum is sagittal in outline and verse plate adjoined to the anterior half of the prolonged medially. The anterior prolongation is pronotal deflection. Posteriorly, the plate is broadly inflected, particularly so laterad to the prosternal short and tumid. The posterior one, the meso- intercoxal process. The prosternal inflection, the thoracic intercoxal process, is long and slender, and cryptosternum of Hlavac (1972:126), supports the is invaginated apically to accommodate the apex prosternal apophysis and extends laterally to the of the metathoracic intercoxal process with which it pleural apophysis. The latter is short, broadly forms a strong but flexible connection. Posteriorly, bilobed distally, and continuous with the trochan- the mesosternum is deeply inflexed, particularly tin. (In the prothorax of the clerid Necrobia rufipes beneath its intercoxal process where it forms a DeGeer, the distal end of the pleural apophysis is nearly imperforated septum. Posteroventrally, the also broad but not bilobed (Tremblay, 1958:98).) mesosterneal inflection adjoins a feebly sclerotized The pronotum is densely clothed by short vertical plate located anteriad to a metasternal transverse setae; such setae line the posterior margin of the ridge. (A comparable area in the cantharid Chan- pronotal postcoxal process. The prosternum is pre- liognatus pennsylvanicus (DeGeer) was interpreted NUMBER 227 13 by Matsuda (1970:213) to consist of "mesospinas- of the clerid genera Epiclines Blanchard and Natalts ternal" and "metapresternal" elements.) Laporte. He noted a substantial difference between the metendosternites of N. laplacii Laporte and Metathorax N. impressus (Spinola). The metathorax (Figures 21, 22, 24) is feebly Wings sclerotized dorsally, strongly sclerotized and boldly convex ventrally. The metanotum is typically com- MESOTHORACIC WINGS.—The elytron (Figures 26, plex, being partitioned into various sclerites, 28, 29) is an oblong protective cover that overlays sutures, and/or other structures correlated with the the thorax and all abdominal segments; its posterior development of flight. A lengthy description of deflection, the apical slope (Figure 29), is acute. the metanotal components is not necessary because The dorsal surface is copiously indented with they have no immediate relevance to the taxonomic coarse favus (or honeycombed-shaped) punctations data herein; but an illustration (Figure 24) of some and has a sparsely microrugose microsculpture of the more important regions might be useful for (note arrow in Figure 307). The latter gives the subsequent comparative studies. The metapleuron punctate and interpunctate surface a glittering (Figures 21, 22) consists primarly of the slender arenose appearance that becomes particularly ap- metepisternum; the metepimeron is largely mem- parent under intensive light. The elytral disc also branous, being feebly sclerotized along the meso- has a dense vesture of short reclinate and longer pleural suture and at the pleural wing processes. vertical setae. The metasternum is trapezoidal, about half as In dorsal view, the elytron is outlined by three long as wide, and is divided into two halves by a margins (Figure 26). Two of these, the humeral midlongitudinal suture (the discriminal line of and sutural, are defined according to their mor- Ferris, 1940:37). Ferris apparently was also the phological origin. The third, the posthumeral, is first to suggest that the beetle metathoracic venter, defined in accord with its position and/or function. commonly called the "metasternum," consists of Below the posthumeral margin is the epipleural subcoxal elements. The discriminal line is also margin (Figure 28) that consists of lateral epipleural well-defined in the clerids Enoclerus sphegeus fold and ventral epipleuron. Fabricius and Necrobia rufipes DeGeer (Campau, The humeral margin is feebly sinuous, the post- 1940:78). humeral margin is linear in basal two-thirds and Internally, the discriminal line is expressed by a acutely arcuate in apical third. The sutural margin bold inflection referred to as the discrimen by is linear and has a prominent sutural ridge that Ferris (1940:36) and the median ridge by Doyen extends from the sutural cleft (as in Figure 27) to (1966:123). The inflection is a keel-like structure the elytral apex. The sutural margin has a sutural that nearly spans the length of the metasternum groove and sutural flange that interlock the reposed and increases in depth posteriorly where it becomes elytra. continuous with the internal ridge of the antecoxal METATHORACIC WINGS.—A membranous wing is suture. Posteriad to this suture is a narrow tapered illustrated in Figure 23. Wing venation character- antecoxal piece ("Katepisternum 3" of Campau, istics of JP. reventazon are not described because 1940:78) whose ventral extremity forms the ventral they are of no consequence in delimitation of coxal condyle. perilypine species and because terminology of clerid To my knowledge a metafurcal sclerite, or meta- wing venation is currently being reevaluated by sternellum, is not distinguishable in the Cleridae; other workers. however, a well-formed Hylecoetoid metendosternite (Crowson, 1944:274) does not vary appreciably Legs in Perilypus. The metendosternite (Figure 25) of P. reventazon consists of a broad laterally arcuate Close apposition of the intrasegmental leg bases, stalk, oval lamina, and feebly arcuate jurcal arms as found in the more successful surface predatory that project anterolaterally. Solervicens (1973a, fig. beetles, is a primary structural adaptation for an 5b; 1973b, figs. 4a,b) studied the metendosternites efficient mechanism of cursorial locomotion. Of 14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MESOSTERNUM MESEPIMERON MITAPUURAL WING PROCESS Mf SOTHORACIC INTERCOXAL PROCESS METATHORACIC INTERCOXAL PROCESS METEPISTERNUM METEPIMERON

MESOPIEURAL SUTURE DISCRIMINAL LINE ANTECOXAl SUTURE

FIRST VISIBLE ABDOMINAL STERNUM

STERNAL DEPRESSION

FIGURES 19-23.—Perilypus reventazon: 19, antennal articles 5 to 8 (arrows indicate microsetae); 20, abdominal sterna n, in, ventral view; 21, mesothorax, metathorax, and abdomen, ventral view; 22, mesothorax, metathorax, and abdomen, lateral view; 23, metathoracic wing. (Sale = I mm) NUMBER 227 15

AXILLARY I AXILLARY 3

LAMINA

HUMERAL MARGIN

.SUTURAl 'RIDGE POSTHUMERAl*- .SUTURAL MARGIN \ MARGIN

FIGURES 24-31.—Perilypus reventazon: 24, metanotum and axillary sclerites; 25, metendosternite; 26, elytron, dorsal view; 28, elytron, cross section; 29, elytron, lateral view. P. orthopleuridus: 27, mesobasal region (arrow indicates sutural cleft). P. limbatus: 30, elytron, lateral view. P. quadrilineatus: 31, elytron, lateral view. (Scales = 1 mm) 16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY secondary importance in this scheme are modifica- of a ninth segment may be present in the form of tions of structures and/or proportions of the leg a feebly sclerotized crescentic plate (the interspicular components, particularly those of the tarsal articles. plate) located beneath the aedeagus and between All of these considerations are to some degree the posterior explanation of the spiculi (Figures 37, pertinent to a discussion of the leg structure of 38). Doyen (1966:138) found a dorsal interspicular clerid beetles which, as a group, are almost exclu- plate in Tenebrio and considered it the tenth sively cursorial predators. tergum. Tanner (1927, pi. 6: figs. 56, 57), in his In P. reventazon the intrasegmental coxae are coverage of female genitalia of Tillinae clerids, separated by a thin intercoxal process, which in identified the membrane between segment 8 and the meso- and metathorax communicate with a the ovipositor as the ninth segment. midlongitudinal extension of the subsequent seg- In general the abdominal dorsum (Figure 22) is mental venter. The procoxa (Figure 32) is conical feebly sclerotized. However, tergal sclerotization and more protrusive than the pyrifonn mesocoxa becomes more intense from tergum 1, which is (Figure 33) and transverse metacoxa (Figure 35). entirely membranous, to tergum 8; the sclerotiza- The pro- and mesotrochanter are triangular and the tion of sterna 3 to 8 is about equivalent. The first metatrochanter is oblong. In general, the femora five terga are glabrous, whereas the sixth and are very similar but the front femur is shorter and seventh have bands of microsetae at their posterior slightly more robust than the rest. The tibiae are margins, which are probably instrumental in the also very similar, but protibia differs by having one folding mechanism of the membranous wings. tibial spur, not two (Figure 32). All tibiae are con- Tergum 8, the pygidium, is vested with long spicuously carinate (the tibial carina) on their reclinate setae, and is subquadrate in males (Figure anterior and posterior fades, and each is particu- 39) and transverse in females (Figure 40). The larly densely setose at its inner apical third. lateral area of the abdomen is broadly membranous. The tarsus, the most variable leg region, con- In more advanced coleopterans it is often diffi- sists of five conspicuous tarsomeres, which differ cult to identify components of the anteroventral with respect to length and pulvillar shape on each region of the abodmen. This region was modified leg (Figure 34). The basitarsus (Figure 33) is always substantially during the posterior extension of the the shortest; the second and fifth tarsomere are metathoracic ventral plate, an adaptation yielding about equal in length; and the third is slightly greater surface area for attachment of flight muscles. longer than the fourth. The emargination of the In general, sternum i is considered absent in most distal border of the pulvillus of each tarsus becomes polyphagan beetles but a vestige of this plate is more pronounced from basitarsus to tarsomere four. present in Tenebrio (Doyen, 1966:135). In Perily- Another structural progression involves abridge- pus, sternum i appears to be entirely suppressed. ment of the first two pulvilli from pro- to meta- The second abdominal sternum (Figure 20) is nar- tarsus. The basitarsal pulvillus, which along with row and feebly sclerotized and is fused with an pulvillus 2 is conspicuously asymmetrical on the anterior depression of the third abdominal sternum metatarsus (note left arrow in Figure 36), is reduced to form the metacoxal cavity. Medially however, and obscured by a narrow setal cluster (note right the two sterna form a keel-like intercoxal process arrow in Figure 36). (Figure 20). Forbes (1925:290) described a similar Protarsal claws are sexually dimorphic with re- development of the abdominal venter in the spect to symmetry. In male specimens the inner anobiid Trypopitys sericeus (Say) (now in the genus claw is conspicuously longer than the outer claw Priobium). (Figure 34). In female specimens the protarsal claws The posterior six sterna, which comprise the are symmetrical as are the claws of the middle and visible abdominal venter, are well-sclerotized, hind legs of both sexes. transverse plates, the lateral and posterior margins of which are narrowly and palely indicated. Sterna Abdomen in to VII each have a pair of sternal depressions (Figure 22). To the inner surface of these are at- The abdomen (Figures 21, 22) is soft, flexible, tached short longitudinal muscles that connect to with eight apparent segments. In males, rudiments the antecostal ridge. Doyen (1966:136) found similar NUMBER 227 17

FIGURES 32-42.—Perilypus reventazon: 32, front leg; 33 middle leg; 34, front tarsus; 35, hind leg; 36, hind tarsus (top arrow indicates pulvillus 2, bottom arrow indicates basitarsai pulvillus); 37, spicular fork, ventral view; 38, spicular fork, lateral view; 39, male pygidium; 40, female pygidium; 41, male abdominal sternum vm; 42, female abdominal sternum vm, (Scales = 1 mm) 18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY muscle attachment regions on exposed sterna of similar glands in the clerid Necrobia ruficollis Tenebrio molitor. The posterior margin of sternum Latreille and indicated that the duct of each gland VII (Figure 21) is broadly emarginated in both sexes, opened on either side of the anus. Scott (1919:108, whereas that of sternum vm is sexually dimorphic, footnote) suggested that the glands reported by being emarginated in males (Figure 41) and arcuate Cholodkovsky might be neotenic structures that in females (Figure 42). The exposed sterna are in mature larvae produce secretions important to vested with long, reclinate setae. construction of the pupal cocoon. During studies of clerid internal anatomy I found the spicular Reproductive Organs gland present in both sexes of species from all suprageneric taxa. In females, the spicular gland MALE.—The aedeagus (sensu Lindroth, 1957:242) connects to the membranous sac of the spiculum consists of two major structures, the tegmen (Figure ventrale (Figure 42). 43) and the phallus (Figure 46). The tegmen is a FEMALE.—The ovipositor (Figures 48, 49), a pre- dorsoventrally compressed cylinder that consists of dominantly membranous tube, connects to the a pair of parameres, the phallobase, two phallobasic eighth segment via an expansive connecting mem- struts, and the phallobasic apodeme. Each paramere brane (the ninth segment of Tanner, 1927, figs. 56, has a basolateral depression, is hollow, and has two 57); three pairs of slender sclerites provide struc- mesal margins (the mesodorsal and the mesoventral tural rigidity to the genital tube. Two pairs of these margins). Together, the parameres enclose a dorsal sclerites, the oblique and ventral bacculi (Figure and ventral sinus (Figure 44). Anteriorly, the pa- 48) are remnants of the first and second valvifers, rameres are continuous with the phallobase, whose respectively (Tanner, 1927:23). The mesoventral anterior limit projects as the mesoventral phallo- extremity of the oblique bacculus articulates with basic apodeme and the lateral phallobasic struts. the posterior limit of the ventral bacculus, which The phallus, which when reposed lies in the at the posterior fourth projects as a short lateral tegmen, consists of one dorsal and one ventral extension (the baccular acumination). The third phallic plate (Figure 46) and two lateral mem- pair of supportive sclerites, proctigeral bacculi (Fig- branes. Each phallic plate is set with a row of ure 49) widen on the proctiger, are fused at pos- marginal denticles aligned along the margin terior third, and are strongly divergent anteriorly. proximal to the phallic plicae. Also, the posterior The coxites (the hemisternites of Lindroth, 1957: ends of these plates fuse to form the phallic apex; 250) are lobate structures that show sclerotized anteriorly each plate projects as a phallic strut. vestiges (the coxital plates) of valvifer two. The The phallic plicae are distended during copulation outer margin of the coxite is broadly vested with (as in Figure 47). Their numerous phallic papillae setae and the inner margin is asetose and strongly (as in Figure 155; top arrow) may be stimulatory sclerotized in the posterior half. Between the dorsal organs or anchorage structures; the latter function and ventral bases of the coxites extend membranous is unlikely because the inner lining of the bursa and acuminate laminae. The dorsal laminal incision copulatrix does not have structures that might be divides the posterior half of the dorsal lamina construed as elements of an interlocking mechanism. (Figure 49) into two equal lobes; this lamina is When reposed the aedeagus rests on the spicular substantially larger than the unilobed ventral fork (Figure 37), which provides an attachment lamina (Figure 48). Posteriorly, the coxite projects surface for the aedeagal protractor and retractor as a well-developed coxital stylus that is multisetose muscles, and guides aedeagal movements. The distally. spicular fork consists of two slender spicules that are broadened posteriorly and fused at the anterior third. Between the spicules and attached to them is INTERNAL MORPHOLOGY the spicular sac, which is opened proximal to the interspicular plate. To the ventrolateral walls of Digestive System this sac connects a spheroid gland (the spicular The male and female alimentary canal of P. gland), whose function has not been determined. reventazon were described by Ekis and Gupta Cholodkovsky (1913:530) described and figured (1971:56, figs. 28, 29, 76, 77) under C. signaticollis NUMBER 227 19

PARAMERE PHALLIC PUCAE ESOVENTRAL MARGIN ESODORSAL MARGIN VENTRAL SINUS PHALLUS DORSAL SINU

PHALLOBASE

TEGMEN

-_PHAUOBASIC STRUTS

43 PHAILOBASIC APODEME 45

COXITAL STYLUS COXITE COXITAL DORSAL LAMII PLATE LAMINAL INCISIO VENTRAL LAMINA OBLIQUE PROCTIGER BACCULUS

BACCULAR ACUMINATION r__VENTRAL ^^ BACCULUS

PROCTIGERAl BACCULUS

FIGURES 45-50.—Perilypus reventazon: 43, tegmen, ventral view, 44, tegmen, dorsal view, 45, tegmen, lateral view, 46, phallus. P. chaletoides: 47, aedeagus, dorsal view; 48, ovipositor, ventral view; 49, ovipositor, dorsal view; 50, brain and ventral nerve cord, dorsal view. (Scales = 1 mm) 20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

OUTER BRANCH

INNER BRANCH

ANTERIOR ACCESSORY GLAND

POSTERIOR ACCESSORY GLAND

SPERMATHECAL CAPSULE PERMATHECAL GLAND DUCT SPERMATHECAL GLAND SPERMATHECAL DUCT 56 BURSA COPULATRIX OVARY MEDIAN OVIDUCT

LATERAL OVIDUCT CALYX

VAGINA

FIGURES 51-56.—Perilypus reventazon: 51, male internal reproductive organs; 52, anterior accessory gland; 53, posterior accessory gland; 54, ejaculatory duct; 55, female internal reproductive organs. P. xtcntralis: 56, hind leg. (Scales = 1 mm) NUMBER 227 21 Spinola and C. bilineatus Gorham, respectively. the bursa copulatrix is correlated with the amount The descriptions and illustrations in that work of semen in its lumen. adequately characterize perilypine digestive organs, The ovaries consist of 12 acrotrophic ovarioles which have no appreciable intrageneric variation. that stem from a well-developed calyx. The latter connects to short, lateral oviducts that communi- Nervous System cate with a narrower more elongate median oviduct. The bursa copulatrix, a saccular extension of the An illustration of the ventral nerve cord (Figure anterior region of the vagina, is recurved basally. 50) is provided to establish a frame of reference for The bursa copulatrix is usually spatulate, but in subsequent intergeneric comparisons. The nerve gravid females it is swollen substantially by seminal cord was not notably varied among the species in fluid. Near the base of the bursa copulatrix the which it was studied. spermathecal capsule, a sclerotized ovoid structure, communicates with the vagina via the spermathecal duct; the latter is distended basally, narrowed and Reproductive Systems convoluted distally. The spermathecal gland con- MALE.—The more important male organs (Fig- nects to the spermathecal capsule via a short brun- ures 51-54) are the testes, vas deferens, accessory neous spermathecal gland duct. The vagina is flexed glands, and ejaculatory duct. near the middle and is tapered. Each of two oblong testes consists of 12 tubular testicular follicles that are encapsulated by the Genus Perilypus peritoneal membrane. The follicles unite with the tubular vas deferens via the vasa efferentia, which Perilypus Spinola, 1841:72 [type-species: Perilypus carbonarius join the ejaculatory duct. An oblong swelling on Spinola, 1844a: 105, by monotypy].—Spinola, 1844a: 103.— Lacordaire, 1857:430.—Desmarest, [1860]:235.—Schenkling, the vas deferens proximal to the ejaculatory duct 1903:13.—Ekis, 1975:26. is the seminal vesicle. To the anterior limit of the Derestenus Chevrolat, 1843:13 [type-species: Derestenus quad- ejaculatory duct there also are attached two pairs rilineatus Chevrolat, 1843:13, by monotypy].—Desmarest, of accessory glands which according to their posi- 1860:238.—Lacordaire, 1857:443.—Gorham, 1878:162.— Cor- tion of insertion are designated as the posterior poraal, 1948:243. accessory glands and the anterior accessory glands. DIAGNOSIS.—The combination of serrate antenna The anterior accessory gland (Figures 51, 52) is (Figure 8) and intricate elytral punctation (Figure very long, undivided, and coiled; its narrower distal 2; punctations are usually honeycomb-shaped) dis- third is tightly convoluted. The inner walls of this tinguish specimens of most species of Perilypus gland has two shallow, longitudinal ridges that from those of other New World genera of Clerinae delimit at least two different secretional regions. (sensu Crowson, 1964). Specimens of the frontalis The products of one of these regions are externally and criocerides groups (involving four species) have visible as a dark band along the inner half of the clubbed antenna. Members of the first group are gland. The posterior accessory gland (Figures 51, 53) superficially similar to some specimens of the genus divides near the base into two variously convoluted Placopterus, but are separated easily from them branches. The outer branch is about one-third (p. 26). Specimens of P. criocerides (Gorham) might longer than the inner branch; the texture and other be confused with those of Systenoderes amaenus visible properties of the secretions of both branches Spinola, which differ by having the elytron boldly are identical. The ejaculatory duct (Figure 54) is convex, sparsely setose, and impunctate. bulbous and sinuous in its anterior half, slender DESCRIPTION.—Form: Variable, commonly rec- and linear in the remainder. tangulate (Figure 2) with elytron plane in basal FEMALE.—The major female organs (Figure 55) two-thirds and deflexed in apical third. Members of are the ovaries, lateral and median oviducts, bursa the quadrilineatus and ornaticollis groups are vari- copulatrix, spermathecal capsule, spermathecal ously oval (Figures 279, 293) and usually compressed gland, and vagina. The shape and external appear- dorsoventrally, whereas those of the chaletoides and ance of these organs are generally homogeneous at viridipennis groups are respectively conspicuously infra- or interspecific level. However, the shape of robust, and short and variously convex. 22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Size: Length 5.4 to 14.4 mm, width 1.6 to 5.2 mm. deflection gradual (Figure 30), very gradual (Figure Females usually more robust than males. 31), or acute (Figure 29). Front tarsal claws boldly Color: Head (including antennae), thoracic (Figure 69) or feebly (Figure 34) asymmetrical, or venter, elytron, legs, and abdomen usually flavo- symmetrical. testaceous or black, or both. Pronotum concolorous, Abdomen: The structures of the abdomen are or lateral regions flavotestaceous to rufous and disc discussed on page 16. Male pygidium varied in size with black vitta; latter percurrent or interrupted and shape, particularly robust in some species at middle; sometimes pronotum predominantly (Figure 217); posterior margin usually entire, rarely flavotestaceous with dark brown or black midapical emarginated (Figure 94). macula. Abdomen usually entirely black in males, Male Genitalia: For a general description, see black and flavotestaceous in females. Elytron con- page 18. Usually strongly pigmented throughout; colorous, fasciate, or vittate (usually vittate or in members of the ornaticollis group phallobase fasciate in females only). marked with two dorsal and two ventral longitudi- Integumental Setae: Head, pronotum, elytron, nal bands. Combined length of phallobase and and legs densely setose; setae particularly long and phallobasic apodeme 1.5 to 10 times longer than copious on antenna and tibia. Antennal setal length paramere. Combined length of paramere and phallo- inversely related to elytral setal length; in lampyri- base 1.0 to 4.8 times longer than phallobasic apo- form specimens (where antenna most serrate) deme. Paramere: well developed; dorsum usually antennal setae exceptionally long, elytral setae convex, explanate medially or not; mesodorsal mar- exceptionally short. gin denticulated or not; venter usually concave; Head: Interocular depressions deeply impressed dorsal and ventral sinuses well defined. Phallus: or shallow, confluent posteriorly or not. Eyes usually marginal denticles present or not; phallic plicae boldly convex; ommatidia fine, as wide as ocular usually well developed. suture. Mouthparts, see pages 9, 11. Antenna ser- Female Genitalia: For a general description see rate (Figure 8); rarely clubbed (Figure 67); articles page 18. Dorsal lamina bilobed (Figure 101), or usually increasing in width from scape to article trilobed (Figure 71), rarely heptalobed (Figure 318); 11; funicular articles filiform (Figure 67) to boldly ventral lamina usually trilobed (Figure 85), rarely serrate (Figure 284). unilobed (Figure 48), or pentalobed (Figure 70); Thorax: For discussion of prothoracic venter, laminae often deeply incised; proctigeral bacculi metathorax, and metathoracic wings and legs, fused (Figure 49) or not (Figure 71); ventral see pp. 9-16. Pronotum usually conspicuously bacculi never fused, acuminate at posterior two- transverse, length rarely equals width; subapical thirds or near middle; coxital plates present or depression deeply impressed throughout, or deeply absent. Internal Organs: The treatise of the alimentary impressed laterally and shallow medially, sinuous canal, ventral nerve cord, and the reproductive or linear; pronotal foveae shallow or deeply im- organs of P. reventazon, (p. 18) adequately serves pressed, punctiform or elongate; side margins of as a generic characterization of the internal organs pronotum proper variously convex; prebasal depres- considered. The accessory glands of the male repro- sion and pronotal collar usually well developed. ductive organs, however, are somewhat variable Mesoscutellum subquadrate, transverse (Figure 312), interspecifically. The most notable variation in- or triangular (Figure 300); rarely stalked (Figure volves the anterior glands, which are feebly coiled 310). Elytron variable; outer margins (epipleural in P. ventralis (Figure 316), and the posterior or posthumeral) broadly arcuate, or straight in glands which are exceptionally long in P. chale- basal two-thirds (or more); arcuate in apical third toides (Figure 277). (or more); surface microsculptured (see arrow in DISTRIBUTION.—The species of this genus occur Figure 307) or not; punctation usually favus, deeply in the New World only. Except for P. ornaticollis, impressed or shallow, and profusely distributed whose northern limit is recorded as Ohio (USA), throughout elytral surface, or punctations small the genus is exclusively Neotropical and ranges and round and restricted to posthumeral region. from Mexico to central Brazil. Most of the species Epipleural fold plane or convex; apical slope occur in Central America, from Mexico to Panama. NUMBER 227 23

Key to the Species of Perilypus

I. Antenna (Figure 67) loosely clubbed, funicular articles usually filiform, rarely serrate 2 Antenna (Figure 8) not clubbed, serrate 3 2.(1) Elytron copiously punctate (Mexico to Nicaragua) {frontalis group) 8 Elytral punctations absent, or few punctations in posthumeral region (Mexico) (criocerides group) 9 3.(1) Elytra (Figures 279, 293) ovate, and/or elytral surface copiously impressed with small shallow punctations; punctate and interpunctate elytral surfaces arenose, rarely smooth and shiny; pronotum (Figure 279) boldly transverse; antenna (Figure 284) usually boldly serrate 4 Elytra (Figure 2) rectangulate; elytral surface impressed with large deeply impressed punctations (Figure 2), punctations profusely distributed (Figure 2), or concentrated in posthumeral region (Figure 99); punctate and interpunctate elytral surface usually smooth and shiny; pronotum (Figure 110) feebly transverse, length rarely equals width; antenna rarely boldly serrate 6 4.(3) Mesoscutellum (Figure 312) transverse; femora usually dark brown or black and with subapical annulus (Figure 56), rarely entirely dark brown or flavotestaceous (USA to Venezuela) (ornaticollis group) 10 Mesoscutellum subquadrate or trigonal; femora predominantly flavotestaceous or entirely dark brown or black, rarely annulated subapically 5 5.(4) Dorsal fades entirely stramineous or predominantly castaneous (Mexico) (chaletoides group) , 17 Dorsal fades not as above; elytral surface entirely black, rarely predominantly piceous and vittate discally (Mexico to Guatemala) (quadrilineattis group) 18 6.(3) Epipleural fold convex, and/or elytral surface impunctate at apical fourth or more 7 Epipleural fold plane, concave or grooved longitudinally; elytral surface coarsely punctate throughout (Mexico to Peru) (reventazon group) 22 7.(6) Elytral apical fourth impunctate or with few very small punctations (but see variation under P. levis) (Panama to Brazil) (viridipennis group) 56 Elytral surface coarsely punctate throughout (Coast Rica to Peru) (limbatus group) 58 8.(2) Front tarsal claws asymmetric; epipleural fold grooved longitudinally (males), or shifted dorsally (females) by boldly convex epipleuron at elytral middle half (Figure 61) (Mexico to El Salvador) 1. P. frontalis (Gorham), new combination Front tarsal claws symmetric; epipleural fold convex from elytral base to elytral apex (El Salvador to Nicaragua) 2. P. setuUis, new spedes 9.(2) Antennal club compact (Figure 80), funicular articles filiform; elytron predominantly flavotestaceous; posterior margin of pygidium feebly emarginated in males (Figure 86), entire in females (Mexico) 3. P. criocerides (Gorham), new combination Antennal club lax (Figure 96), funicular articles serrate; posterior margin of pygidium deeply emarginated in males (Figure 94), feebly emarginated in females (Figure 95); elytron dark blue or purpurescent (Mexico) 4. P. insectus, new spedes 10.(4) Femora entirely dark brown or entirely flavotestaceous, not annulated 11 Femora predominantly black, annulated (Figure 56) 12 11.(10) Pronotal discal vitta furcate (Figures 322-324); elytral discal vitta (when present) not strongly diverged from elytral suture at elytral apical fifth; femora entirely dark brown (USA to Mexico) 43. P. ornaticolUs (LeConte), new combination Pronotal discal vitta reduced to a midbasal macula (Figure 342); elytral discal vitta strongly diverged from elytral suture at elytral apical fifth (Figure 342); femora entirely flavotestaceous (Panama) 49. P. floralis (Gorham), new combination 12.(10) Fourth and fifth visible abdominal sternum flavous; mesoscutellar disc concave; average size about 13 mm (Mexico to Guatemala) 42. P. vtntralu (Gorham), new combination Fourth and fifth visible sternum black; mesoscutellar disc plane; average size from 6 to 8 mm IS 13.(12) Epipleural fold plane 14 Epipleural fold obliquely convex 16 24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

14.(13) Elytron sinuate apically (Figure 354), surface with stout vertical setae, sutural margin diverged at apical fifth (Figure 354) (Panama) 46. P. pilatus, new species Elytron truncate apically, vertical setae and sutural margin normal 15 15.(14) Antennal article 11 as long as preceding three combined (Figure 331) (Mexico) 45. P. prolixicornis, new species Antennal article 11 as long as preceding two combined (Figure 332) (Mexico to Guate- mala) 44. P. galbeus, new species 16.(12) Elytral apex uplifted (Figure 353), surface dull, epipleural fold obliquely convex (Costa Rica) 47. P. tmtarius, new species Elytral apex not uplifted but sinuatotruncate. surface shiny, epipleural fold evenly convex (Venezuela) 48. P. sinuapicis, new species 17.(5) Elytral surface entirely stramineous (Mexico) 36. P. bicolor (Chevrolat), new combination Elytral surface predominantly rufous, apex and sutural margin usually black (Figure 266), rarely only faintly infuscated at suture (Mexico) 35. P. chaletoides, new species 18.(5) Epipleural fold obliquely convex, visible in specimen dorsal view; elytral vitta as in Figure 279 (Mexico to Guatemala) 37. P. quadrittneatus (Chevrolat), new combination Epipleural fold not obliquely convex and not visible in specimen dorsal view; elytral disc avittate 19 19.(18) Elytral surface undulated longitudinally (Figure 305); pronotum not coarsely macrosculptured 20 Elytral surface not undulated; pronotum coarsely macrosculptured 21 20.(19) Pronotal vitta exceptionally broad, with oblong roseous macula at center; femur testaceous ventroapically (Mexico) 39. P. coronijormis, new species Pronotal vitta constricted medially, not maculated centrally, femora annulated subapically (Guatemala) 40. P. fiuctus, new species 21.(19) Elytral disc bicarinate (Figure 300); pronotum concave centrally and paralaterally. coarsely punctate (Mexico) 41. P. bicristatus, new species Elytral disc not carinate; pronotum plane, transversely wrinkled; posthumeral margin briefly vitiate (Figure 287) (Mexico to Guatemala) 38. P. telephoroides (Gorham), new combination 22.(6) Posterior margin of sixth visible sternum emarginated (Figure 41) (males) 23 Posterior margin of sixth visible sternum not emarginated (Figure 42) (females) 39 23.(22) Front tarsal claws boldly asymmetrical 24 Front tarsal claws feebly asymmetrical, or symmetrical 28 24.(23) Length about 6 mm; elytron olive green or blue green, apex usually testaceous (Mexico) 32. P. immitis, new species Length more than 8 mm; elytral coloration not as above 25 25.(24) Antenna testaceous (Mexico to Costa Rica) 27. P. orthoplettridus (Thomson), new combination Antenna entirely black or dark brown 26 26.(25) Pygidium transverse (Mexico) 29. P. latilobus, new species Pygidium oblong 27 27.(26) Antennal articles 3-5 moderately serrate (Figure 8); procoxae infuscated (Guatemala) .... 26. P. calcaris, new species Antennal articles 3-5 feebly serrate, nearly filiform; procoxae immaculate (Mexico) ... 28. P. exttis, new species 28.(23) Elytral apex testaceous (Panama) 23. P. bilineatus (Gorham) Elytral apex not testaceous 29 29.(28) Antenna boldly serrated (Figure 244) SO Antenna moderately serrated 31 30.(20) Femora infuscated at apex only; elytra not truncated (Panama) 24. P. cerroaxul, new species Femora infuscated at distal third or more; elytra truncated (Peru) 31. P. crassut, new species 31.(29) Specimens north of Nicaragua 32 Specimens south of Nicaragua 35 NUMBER 227 25

32.(31) Body form slender (Figure 239); elytra feebly convex; elytral vertical setae about one- third longer than elytral reclinate setae; pronotal vitta constricted at subapical depression (Mexico to Guatemala) 30. P. distinctvu (Chevrolat), new combination Body form squat; elytra plane; elytral vertical setae about one-half longer than elytral reclinate setae; pronotal vitta not as above S3 33.(32) Pronotal subapical depression shallow at middle; pronotal arch feebly convex, ap- proaching plane (Guatemala) 19. P. nigriventris (Gorham) Pronotal subapical depression deeply impressed throughout; pronotal arch boldly convex 34 34.(33) Parameres short, strongly arcuate (Figure 255); parameres with combined venter deeply concave (Figure 255) (Mexico) 33. P. caliculus, new species Parameres not as above, but as in Figure 258 (Mexico) 34. P. carbonarius Spinola 35.(31) Profemur black in ventroapical third or entirely flavotestaceous (Costa Rica) 21. P. claudut (Wolcott) Profemur flavotestaceous in ventroapical third 36 36.(35) Metafemur entirely flavotestaceous or infuscated dorsoapically S7 Metafemur black in apical half or less (Costa Rica) 20. P. sapient is, new species 37.(36) Antenna more serrate than in Figure 8; metafemur flavotestaceous (Costa Rica) 22. P. cultratus, new species Antennal serration as in Figure 8 38 38.(37) Metafemur infuscated dorsopreapically (infuscation narrowed to femoral apex dorso laterally or not narrowed) (Costa Rica) 18. P. rrventazon, new species Metafemur infuscated dorsoapically (Panama) 25. P. ordinatus, new species 39.(22) Epipleuron broadly explanate in elytral middle third (Figure 230) (Mexico) 27. P. orthopUuridus (Thomson) Epipleuron not broadly explanate 40 40.(39) Sixth visible abdominal sternum conspicuously arenose and with broad shallow depressions; fourth and fifth visible sternum flavous (Guatemala) 26. P. calcaris, new species Abdominal sterna not as above 41 41.(40) Elytral disc vittate (sometimes vitta only faintly visible) 42 Elytral disc avittate 51 42.(41) Elytral discal vitta closer to posthumeral margin than to sutural margin (Figure 186) 43 Elytral discal vitta about equidistant from side margins (Figure 193) 46 43.(42) Elytral discal vitta gradually incurved posteriorly, terminating at elytral apex 44 Elytral discal vitta acutely incurved posteriorly, terminating at sutural margin before elytral apex 45 44.(43) Elytral sutural margin flavotestaceous at basal third or more (Mexico) 34. P. carbonarius Spinola Elytral sutural margin piceous (Panama) ... 23. P. bilineatus (Gorham), new combination 45.(43) Femora black in distal half or more, not flavotestaceous ventroapically (Costa Rica) 21. P. claudus (Wolcott) Femora infuscated in dorsodistal half, flavotestaceous ventroapically (Costa Rica) 18. P. reventaum, new species 46.(42) Anterior limit of elytral discal vitta before humeral margin (Figure 239); first five visible abdominal sterna flavotestaceous 47 Anterior limit of elytral discal vitta at humeral margin (Figure 186); abdominal sternum not as above 48 47.(46) Elytron feebly convex; metafemur infuscated at dorsoapical third (Mexico) 30. P. distinct™ (Chevrolat) Elytron plane; metafemur infuscated at apex only (Costa Rica) 22. P. cultratus, new species 48.(46) Femora predominantly black (Mexico) 29. P. latilobus, new species Femora predominantly flavotestaceous 49 49.(48) Antenna flavotestaceous (Costa Rica) 21. P. claudm (Wolcott) Antenna black 50 50.(49) Meso- and metafemur black in apical third or more (Costa Rica) 20. P. sapientis, new species 26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Meso- and metafemur black at apex only (Guatemala) 19. P. nigriuentris (Gorham), new combination 51.(41) Elytron broadly flavotestaceous at or near margin, fasciate (Figures 262-265), or entirely Bavotestaceous (Mexico) 34. P. carbonarhu Spinola Elytron not as above, disc entirely black 52 52.(51) Antenna boldly serrate (Figure 244); elytra feebly convex (Peru) 31. P. crassus, new species Antenna moderately serrate (Figure 8); elytra plane 53 53.(52) Elytral apex testaceous and/or elytra olive green or blue green (Mexico) 32. P. unmitu, new species Elytral apex not testaceous 54 54.(53) Femora flavotestaceous in basal third; pronotal vitta abruptly narrowed at subapical depression (Mexico) 29. P. latUobus, new species Femora entirely black; pronotal vitta not as above 55 55.(54) Pronotum boldly transverse (Mexico) 34. P. carbonarhu Spinola Pronotum feebly transverse (Mexico) 33. P. caliculut, new species 56.(7) Antenna boldly serrate (Figure 244) and densely setose (Canal Zone to Brazil) 7. P. levis, new species Antenna only moderately serrate and moderately setose 57 57.(56) Antenna uniformly black; elytral apical fourth impunctate or with very small punctations; epipleural fold more collapsed than convex (Panama to Colombia) 6. P. relucens (Gorham), new combination Antenna predominantly dark brown, flavotestaceous apically; elytral apical half impunctate; epipleural fold convex (Panama to Venezuela) 5. P. viridipennis (Spinola), new combination 58.(7) Legs testaceous, except hind femur black in distal half (Panama) 16. P. tettaceicornis (Pic), new combination Combination of leg colors not as above 59 59.(58) Epipleural fold wider than antennal pedicel (north of Colombia) 60 Epipleural fold not wider than antennal pedicel (South America) 62 60.(59) Elytral apex black; elytral discal vitta sharply incurved and broadly outcurved posteriorly (Figure 2) (Panama) 17. P. decoris, new species Elytral apex flavotestaceous; elytral disc avittate, or vitta not curvate posteriorly 61 61.(60) Specimens from north of Panama; male genitalia as in Figures 172-174 15. P. prolixipenis, new species Specimens from Panama; male genitalia as in Figures 167-169 14. P. latilira, new species 62.(59) Distal half of hind femur predominantly or entirely black; frons partially or totally black (Figures 58, 59) 63 Distal half of hind femur predominantly flavotestaceous, only apex infuscated; frons flavotestaceous (Figure 57) 66 63.(62) Inner margins and venter of antennal articles 9 to 11 testaceous (Colombia) 10. P. apocopates, new species Inner margins and venter of antennal articles 9 to 11 black 64 64.(63) Femur entirely black 11. P. Ms, new species Femur flavotestaceous basally 65 65.(64) Labrum and frons black (Figure 59) (Colombia) 9. P. cohmtbicus (Spinola), new combination Labrum testaceous; frons testaceous anteriorly, black posteriorly (Figure 58) (Colombia) 13. P. buga, new species 66.(62) Antennal articles 9 to 11 testaceous ventrally (Venezuela to Bolivia) 8. P. Umbatus (Gorham), new combination Antennal articles 9 to 11 black ventrally (Peru) 12. P. acus, new species

The frantalis Group other species of Perilypus except the two species that constitute the criocerides group. Specimens of The members of this group have clubbed anten- the frontalis group differ from those of the crio- nae, a characteristic that separates them from all cerides group by having the elytral surface copiously NUMBER 227 27

FIGURES 57-61.—Anterior limit of black color of frons: 57, Perilypus limbatus, 58, P. buga, 59, P. columbicus. P. frontalis: 60, antenna, dorsal view, X 60 (arrows indicate apical border of antennal articles); 61, female elytron, lateral view, X 27 (arrow indicates convex epipleuron). (Scale = 1 nun) 28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY and deeply punctated; the paramere (Figures 73, macula, or clypeus, frons, and cranial venter flavo- 78) long, feebly pigmented, and without lateral testaceous and epicranium and gena black; antenna depression; and the phallic plate (Figures 74, 79) black; prothorax either flavotestaceous at sides and devoid of marginal denticles. with percurrent or medially interrupted black discal Specimens of the frontalis group superficially vitta, or flavotestaceous or pale orange and with or resemble those of the genus Placopterus (Wolcott, without dark brown midapical macula and dark 1910b: 363) from which they can easily be distin- brown collar; femur either flavotestaceous in basal guished as follows: in frontalis group specimens the four-fifths and black in apical fifth, flavotestaceous antenna extends beyond the pronotal collar, anten- in basal fifth and black in apical four-fifths, or nal article 11 is oblong, body form is rectangulate, entirely flavotestaceous; tibia black or yellow; meso- and pronotal subapical depression is sinuous. In and metasternum piceous; elytron either shiny Placopterus species the antenna does not reach the black but apex flavotestaceous (Figure 62), black pronotal collar, antennal article 11 is quadrate or with sutural and marginal flavotestaceous vitta transverse, body form is more oval, and the pronotal (Figure 63), as in Figure 64, or stramineous and subapical depression is broadly arcuate. infuscated subapically (Figure 66); abdomen black. The distribution of this group extends from Head: Interocular depressions broad and shal- Chiapas, Mexico, to Rivas, Nicaragua. low; eyes shallow; frons (Figure 65) conspicuously broad (HW/IOW, average about 1.9, range 1.8-2-1; 10 males and 10 females measured); antenna 1. Perilypus frontalis (Gorham), new combination clubbed (Figures 60, 67); funicular articles feebly serrate; apical border and disc of antennal articles 7 FIGURES 60-76, 356 and 8 not microsetiferous; AL/PLS, about 1.7; Poecilochroa frontalis Gorham, 1886:338 [holotype: female, length /width ratio of each male antennal article deposited in BMNH; type-locality: Ciudad Guatemala, 2.5:1.4:2.0:2.0:2.0:1.7:1.8:1.7:1.3:1.1:1.7. Departamento de Guatemala, Guatemala]. Pronotum: PL/PW, average about 0.86, range Derestenus apicalis Pic, 1941:10 [lectotype: female, deposited in MNHP, here designated; type-locality: Guatemala; new 7.9-9.2 (10 males and 10 females measured); sub- synonymy]. apical and prebasal depressions deeply impressed; side margin of pronotal proper boldly convex. Pic's type-specimen does not differ significantly Mesoscutellum: Subquadrate. from Gorham's type-specimen. Elytron: Epipleural margin straight in basal DIAGNOSIS.—The extraordinary degree of asym- three-fourths, broadly arcuate in apical fourth; slope metry between the inner and outer front tarsal of apical deflection gradual; discal punctations well claws (Figure 68) will distinguish males of this impressed and evenly distributed; punctate and species from those of any other congeneric species. interpunctate surfaces smooth, shiny, and densely Female specimens are easily recognized by the covered with pale reclinate setae; vertical setae boldly convex epipleuron, which is most developed piceous; epipleuron and epipleural fold variable in elytral middle half. The modification of the (see "Variation"); EL/EW, average about 4.3, range female epipleuron causes the epipleural fold to be 3.9-4.9 (10 males and 10 females measured). shifted upward and the confluence of both struc- Front Tarsal Claws: Strongly asymmetrical in tures gives the epipleural margin a tubular appear- both sexes; inner claw abom *— e times longer ance (Figure 61). than outer claw in males (tifc-.e 69), and about DESCRIPTION.—Form: As in Figure 62. one and a half times longer in females (Figure 68). Size: Length: males, average about 6.3 mm, range Male Genitalia (Figures 72-74): Feebly pig- 5.2-7.4 mm; females, average about 6.7 mm, range mented. Phallobase very broad. Combined length 5.7-7.8 mm. Width: males, average about 1.9 mm, of phallobase and phallobasic apodeme 1.5 times range 1.8-2.3 mm; females, average about 2.2 mm, longer than paramere. Combined length of para- range 1.8-2.7 mm. Ten males and 10 females mere and phallobase 3.0 times longer than phallo- measured. basic apodeme. Paramere: apex obtuse, feebly Color: Cranium either flavotestaceous, pale declinate in lateral view; lateral depression absent; orange, flavotestaceous with midepicranial black dorsum convex, explanate medially; venter narrow, NUMBER 227 29

62 63

FIGUKES 62-66.—Perilypus frontalis: 62-63, habitus; 64, elytra; 65, head, front view, X 30; 66, elytra. (Scale = I mm) 30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY mesoventral margin broadly arcuate; mesodorsal LOCALITY RECORDS (Figure 356).—I examined 110 adult margin sinuous. Sinus: dorsal narrow, lanceolate; specimens from Central America. MEXICO: Estado de Chi- apas: Chiapas (ZMAN, 1 female) ; 2 miles [3.2 km] north- ventral elliptical. Phallus: marginal denticles ab- west of Pueblo Nuevo (GHNE, 1 female); 20 miles [32 km] sent; phallic plicae well developed (47 specimens north of Bochil, Yerba Buena (CNCC, 1 female; GEKI, 2 examined). females); San Crist6bal de las Casas (GEKI, 1 male and Female Genitalia (Figures 70, 71): Laminae very 1 female); Huistan area, 27 miles [43 km] northeast of San Cristobal de las Casas (GEKI, 1 male); 11 miles [18 km] short; dorsal lamina trilobed, lobes equal in length; northeast of San Cristobal de las Casas (GEKI, 1 female); ventral lamina pentalobed, paralateral pair of lobes 7 miles [11 km] southeast of San Cristobal de las Casas shorter than medial lobe; proctigeral bacculi widely (HFHO, 1 male); Teopisca (MNHB, 1 male and 5 fe- separated; ventral bacculus acuminate at posterior males; GEKI, 2 males and 5 females); Highway 24, 8 to 10 two-thirds; coxital plates absent (2 specimens miles [13 to 16 km] southeast of Teopisca (CNCC, 4 males and 3 females; GEKI, 2 males and 2 females; HFHO, 1 examined). male and 3 females) ; Comitan (GEKI, 7 males and 5 fe- Internal Reproductive Organs: Male (Figures 75, males; MNHB, 7 males and 13 females); El Rincon, route 76); anterior accessory gland uniramous, tightly 17 (HFHO, 1 female); Chincultic, near El Rincon (GEKI, coiled; posterior accessory gland biramous, outer 1 female); Zinacatan (GEKI, 1 male and 2 females; NMNH, branch little less than twice as long as inner branch; 1 female). GUATEMALA: Departamento de Guatemala: testis composed of 12 follicles. Female, as in Figure Ciudad Guatemala (BMNH, 1 female, holotype); Departa- mento de Sacatepequez: Duerias (BMNH, 1 male and 1 fe- 55 (5 males and 2 females examined). male; GEKI, 1 female); Capetillo (BMNH, 2 males and 1 VARIATION.—Structural: The degree of asym- female; GEKI, 4 males and 1 female; MNHB, 1 male; MNHP, metry of the front tarsal claws varies with sex (see 2 females; NMNH, 1 male). Departamento de Chimalte- "Front Tarsal Claws") as does development of the nango: Calderas (BMNH, 1 female; GEKI, 1 female). epipleuron and the epipleural fold. In the female "Guatemala" (MNHP, 1 female). EL SALVADOR: De- partamento de Santa Ana: 23 kilometers north of Metapan elytron the epipleuron and epipleural fold are (GEKI, 1 female). Departamento de San Miguel: Cerro boldly convex (Figure 61); in males the epipleuron Verde (CASC, 1 male; GEKI, 4 males and 2 females; HFHO, is not convex and the epipleural fold is grooved 6 males and 2 females). longitudinally. The degree of asymmetry of the front tarsal claws 2. Perilypus sensilis, new species also varies slightly independent of sex. The epipleuron and epipleural fold is collapsed in some female FIGURES 77-79, 356 specimens; however, this variation is probably due TYPE-LOCALITY.—Cerro San Jer6nimo, Departa- to incomplete cuticular expansion. mento de La Paz, El Salvador. Color: Variation in color, not correlated with TYPE-SPECIMENS.—The holotype male is de- sex or geography, is extreme (Figures 62-66). In posited in MNHP, the allotype in BMNH. Both general, color of various parts of the integument specimens were collected by G. C. Champion. Two not normally black or brown ranges from stramine- paratypes: GEKI, 1 male: HFHO, 1 male. ous to fulgidus. DIAGNOSIS.—Distinguishable from P. frontalis, NATURAL HISTORY.—In June, in Sumpango, the only other member of the frontalis group, by Guatemala, I collected four specimens at 2012 symmetry of the front tarsal claws, by the evenly meters and in Zinacatan, Mexico, three specimens at convex epipleural fold (male and female), and by 1830 meters. The Guatemalan sample was collected the less convex side margins of the pronotum by beating roadside shrubs while those from Mexico proper. were collected by beating oak branches (Quercus DESCRIPTION.—Form: As in P. frontalis (Figure sp.). In May, on Cerro Verde, El Salvador, H. F. 62). Howden collected 13 specimens at 2000 meters. Size: Length: males, average about 6.8 mm, range Adult specimens were also collected in July, in 5.6-8.0 mm; females 7.6 mm. Width: males, aver- Mexico. age about 2.2 mm, range 1.8-2.9 mm; females 2.2 DISTRIBUTION (Figure 356).—The known range mm. Three males and 1 female measured. of this species extends from southern Mexico to Color: Clypeus, from, and cranial venter flavo- southern £1 Salvador and southwestern Guatemala. testaceous, epicranium and gena black; antenna NUMBER 227 31

FIGURES 67-74.—Perilypus frontalis: 67, antenna; 68, female front tarsal claw, X 140; 69, male front tarsal claw, X 140; 70, ovipositor, ventral view; 71, ovipositor, dorsal view; 72, tegmen. lateral view; 73, tegmen, ventral view; 74, phallus. (Scale = 1 mm) 32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICURFS 75-80.—Perilypus frontalis: 75, male internal reproductive organs; 76, posterior accessory glands. P. sensilis: 77, tegmen, lateral view; 78, tegmen, ventral view; 79, phallus. P. criocerides: 80, antenna. (Scales = 1 mm) NUMBER 227 black; prosternum and side margin of pronotum LOCALITY RECORDS (Figure 356).—I examined 4 adult flavotestaceous; pronotal disc broadly vittate; specimens from Central America. GUATEMALA: Depart- femora flavotestaceous in basal four-fifths, dark amento de Alta Verapaz: Purulha (BMNH, 1 female). EL SALVADOR: Departamento de San Salvador: Boquer6n, near brown in apical fifth; tibia and tarsus dark brown; Santa Tecla (HFHO, 1 male) ; Departamento de La Paz: meso- and metasternum black; elytron flavotestace- Cerro San Jeronimo (MNHP, 1 male, holotype). NICA- ous along epipleural margin and at apex, black RAGUA: Departamento de Rivas: San Juan del Sur (GEK.I, in remainder; abdomen black. 1 male). Head: Cranium and antenna as in P. frontalis except eyes more convex and AL/PLS 1.9. The criocerides Group Pronotum: As in P. frontalis except less transverse (PL/PW, average about 0.93, range 0.87-0.98; The group consists of two rather dissimilar spe- 4 specimens measured) and side margin of pro- cies. In both of these the antenna is clubbed, notum proper less convex. elytral surface smooth and shiny and only sparcely Elytron: As in P. frontalis except surface less vested with small punctations behind the humerus, shiny, punctations more conspicuous, and epi- and the male pygidium is conspicuously emargin- pleural fold (of both sexes) evenly convex from ated posteriorly (Figures 89, 94). Components of the elytral base to elytral apex. male genitalia are strongly pigmented and phallic Front Tarsal Claws: Symmetrical in both sexes. marginal denticles are present. In P. criocerides the Male Genitalia (Figures 77-79): Feebly pig- phallic plates are denticulated on both margins mented. Phallobase very broad. Combined length (Figure 84), a characteristic not present elsewhere in of phallobase and phallobasic apodeme 3 times Perilypus. longer than paramere. Combined length of para- Members of this group are known only from mere and phallobase 2.8 times longer than phallo- southern Mexico. basic apodeme. Peramere: apex broadly incurved and obtuse; apical third declinate in lateral view; 3. Perilypus criocerides (Gorham), lateral depressions absent; dorsum convex, broadly new combination explanate medially; venter narrow and feebly concave; mesoventral margin broadly concave; meso- FIGURES 80-87, 356 dorsal margin sinuous. Sinus: dorsal narrowly Colyphus criocerides Gorham, 1882:144 [holotype: male, de- lanceolate; ventral elliptical. Phallus: as in P. posited in BMNH; type-locality: Cerro de Plumas, Estado frontalis except phallic plates broader (3 speci- de Veracruz, Mexico]. mens examined). Female Genitalia: Not studied. DIAGNOSIS.—The compact yellow antennal club VARIATION.—Structural: No structural variation (Figure 80) and nearly impunctate and predomi- was noted. nantly flavotestaceous elytron (Figure 81) will dis- Color: The only noticeable variation is in the tinguish members of this species from other species intensity of the vitta on the elytral epipleural mar- occurring in Mexico. gin; the vitta is most pronounced in specimens DESCRIPTION.—Form: As in Figure 81. from Guatemala. Size: Length: male, 8.0 mm; females, average NATURAL HISTORY.—G. C. Champion collected about 7.7 mm, range 7.4-8.1 mm. Width: male 2.2 the holotype from Cerro San Jer6nimo, Guatemala, mm; females, average about 4.4 mm, range 4.3-4.5 at 914 meters. In May, in Boquer6n, El Salvador, mm. One male and 3 females measured. H. F. Howden collected one specimen at 1800 Color: Cranium flavotestaceous; first 8 antennal meters. articles dark brown, article 9 to 11 flavotestaceous; DISTRIBUTION (Figure 356).—The known range prothorax flavotestaceous, pronotum with or with- extends from Alta Verapaz, Guatemala, to San Juan out midapical macula or percurrent discal vitta; del Sur, Nicaragua. meso- and metasternum black; femur predomi- ETYMOLOGY.—Latin, adjective sensilis (sensitive), nantly flavotestaceous, infuscated apically; tibia and with reference to the inferred sensory function of tarsus dark brown; elytron entirely flavotestaceous, the microsetae on antennal articles 9 to 11. or predominantly flavotestaceous and with one 34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY basosutural and one discal black macula (Figure subapical depression deeply impressed and strongly 81); abdomen dark brown. sinuous, side margin of pronotum proper boldly Head: Frons broad (HW/IOW, 1.9; 4 specimens convex. measured); interocular depressions broad, deeply Mesoscutellum: Subquadrate. impressed; eyes boldly convex; antenna (Figure 80) Elytron: Posthumeral margin straight in basal distinctly clubbed, funicular articles filiform, arti- half, broadly arcuate in apical half; slope of apical cles 7 and 8 without microsetae; length/width ratio deflection gradual; surface punctations faintly visi- of each male antennal article 2.0:1.0:2.2:2.2:2.2: ble posthumerally and near epipleural margin; 2.0:1.6:1.4:0.85:0.60:1.1. punctate and interpunctate surfaces smooth and Pronotum: Feebly transverse (PL/PW, average shiny; epipleural fold grooved longitudinally proxi- about 0.91, range 0.88-0.93; 4 specimens measured), mal to epipleural margin, feebly convex more dor-

FIGURFS 81-87.—Penllypus criocerides: 81, habitus; 82, tegmen, lateral view; 83, tegmen, ventral view; 84, phallus; 85, ovipositor, ventral view; 86, male pygidium; 87, ovipositor, dorsal view. (Scales = 1 mm) NUMBER 227 35 sad; EL/EW, average about 4.4, range 4.3-4.5 (4 DESCRIPTION.—Form: As in Figure 88. specimens measured). Size: Length: male, average about 6.0 mm, range Abdomen: Posterior margin of pygidium emar- 5.8-6.2 mm; female, 6.0 mm. Width: male, average ginated in males (Figure 86), arcuate in females. about 1.8 mm, range 1.7-1.9 mm; female 1.8 mm. Front Tarsal Claws: Asymmetrical in both sexes. Two males and 1 female measured. Male Genitalia (Figures 82-84): Strongly pig- Color: Cranium flavotestaceous; antenna flavo- mented. Combined length of phallobase and phal- testaceous to light brown; prothorax flavotestaceous, lobasic apodeme 5 times longer than paramere. except pronotum with brown midapical macula; Combined length of paramere and phallobase 2.7 mesoscutellum flavotestaceous; pro- and mesofemur times longer than phallobasic apodeme. Paramere: flavotestaceous, except infuscated dorsoapically; lateral depression present; dorsum feebly convex; metafemur flavotestaceous in basal half, brown in venter feebly concave; mesoventral margin concave; apical half; tibia and tarsus brown; mesosternum mesodorsal margin irregular, feebly explanate flavotestaceous, metasternum brown; elytron shiny, medially, and boldly braced basally. Sinus: dorsal metallic dark blue or purple; abdomen black. lanceolate; ventral broadly elliptical. Phallus: Phal- Head: As in P. criocerides except frons broader; lic plates denticulated on both margins; phallic HW/IOW, 1.7 (3 specimens measured); antennal plicae well developed (1 specimen examined). club less compact, funicular articles serrate (Figure Female Genitalia (Figures 85, 87): Dorsal and 96); and length/width ratio of each male antennal ventral lamina trilobed, lateral lobes shorter than article 2.1:1.3:1.4:1.7:1.3:1.7:1.4:1.4:1.4:1.2:2.0. median lobe; proctigeral bacculi widely separated; Pronotum: As in P. criocerides except side mar- ventral bacculus acuminate near posterior two- gins of pronotum proper less convex. thirds; coxital plates absent (1 specimen examined). Elytron: As in P. criocerides except basal half be- VARIATION.—Structural: The available specimens hind humerus finely punctate and distal half im- are structurally homogeneous. punctate. Color: The dorsum of one female is entirely Abdomen: Posterior margin of pygidium deeply flavotestaceous. The black markings on the prono- emarginated in male (Figure 94), feebly emargin- tum and the elytron vary in size. ated in female (Figure 95). NATURAL HISTORY.—J. M. Campbell collected Front Tarsal Claws: Asymmetrical in both sexes. three adults during July, in Chiapas, Mexico. Male Genitalia (Figures 89-91): Strongly pig- DISTRIBUTION (Figure 356).—Known only from mented. Combined length of phallobase and phal- Veracruz and Chiapas, Mexico. lobasic apodeme 3.5 times longer than paramere. LOCALITY RECORDS (Figure 356).—I examined 4 adult Combined length of paramere and phallobase 2.6 specimens from Central America. MEXICO: F.stado de times longer than phallobasic apodeme. Paramere: Chiapas: 17 kilometers north of Tuxtla Gutierrez (GEKI, 3 females) . Estado de Veracruz: Cerro de Plumas (BMNH, carinate lateroapically; attenuate; lateral depression 1 male, holotype). well impressed and localized medially; dorsum feebly convex; venter concave; mesoventral margin 4. Perilypus insectus, new species strongly sinuous; mesodorsal margin feebly sinuous and boldly braced basally. Sinus: dorsal subparallel, FICURES 8&-96, 356 ventral broadly elliptical. Phallus: phallic plate TYPE LOCALITY.—Mexico. broad to phallic plicae; marginal denticles extended TYPE-SPECIMENS.—The holotype male and the from basal third to phallic plicae (2 specimens ex- allotype, deposited in MNHB, were collected by J. amined). Flohr G. One paratype: GEKI, 1 male. Female Genitalia (Figures 92, 93): Laminae and DIAGNOSIS.—The extent of emargination of the coxites very short; dorsal lamina trilobed, lobes pygidial posterior margin (Figure 94) distinguishes about equal in length; ventral lamina pentalobed, male specimens. In female specimens the pygidium paralateral pair of lobes shorter than medial lobe; is feebly emarginated (Figure 95). Small size and proctigeral bacculi widely separated; coxital plates metallic luster of the elytron are diagnostic char- absent. acteristics for both sexes. VARIATION.—Structural: Emargination of the py- 36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 88-95.—Perilypus insectus: 88. habitus; 89, tegmen, lateral view; 90, tegmen, ventral view; 91, phallus; 92, ovipositor, ventral view; 93, ovipositor, dorsal view; 94, male pygidium; 95, female pygidium. (Scales = 1 mm)

gidium is sex dimorphic (Figures 94, 95) as is the DISTRIBUTION (Figure 356).—Known only from degree of asymmetry of the front tarsal claws; the "Mexico." claws are more asymmetrical in males than in the LOCALITY RECORDS (Figure 356).—I examined 3 adult female. specimens from Central America. "Mexico" (GEKI, 1 male; Color: Elytron may be dark blue or purple. The MNHB, 1 male, holotype, and 1 female). antenna varies in extent of paleness. The vtridipennis Group ETYMOLOGY.—Latin, insectus (to cut into; masculine past participle of inseco). I refer to the emar- The conspicuous absence of deeply impressed gination of the pygidial posterior margin. punctations from the apical third (or more) of the NUMBER 227 37 elytron is the major diagnostic characteristic for the equals width (PL/PW, average about 0.95, range group. Others are antenna pronouncely serrate, and 0.90-1.0; 10 males and 10 females measured); sub- elytral punctate and interpunctate surface smooth apical and prebasal depressions deeply impressed; and shiny. Elytral coloration (Figures 103, 110) is side margin of pronotum proper moderately arcu- unique in Perilypus. ate. Geographically, the members of this group are Elytron: Epipleural margin straight in basal half, distributed from northern Panama to northeastern broadly arcuate in apical half; elytral convexity Brazil with distribution records from northern varied (see "Variation); large punctations localized Colombia and Venezuela. behind humerus (Figure 99); punctate and interpunctate surfaces smooth and shiny, not arenose; 5. Perilypus viridipennis (Spinola), epipleural fold feebly convex, but lower region new combination slightly collapsed; slope of apical deflection gradual; EL/EW, average about 4.2, range 3.8-5.0 (20 speci- FIGURES 97-108, 357, 358 mens measured). Front Tarsal Claws: Male, feebly asymmetrical. Systenoderes viridipennis Spinola 1844a: 132 [holotype: sex not determined, deposited in MSPI; type-locality: Colom- Female, symmetrical. bia].— Desmarest, [1860]:239.—Ekis, 1975:27. Male Genitalia (Figures 104-106): Strongly pig- Cleronomus amaenus Schenkling, 1906:263 [lectotype: male, mented. Combined length of phallobase and phal- deposited in DEIE, and 1 paralectotype, female, deposited lobasic apodeme 6 times longer than paramere. in ZMAN, here designated; type-locality: "Magdalen- Combined length of paramere and phallobase 4.0 strom" = Rio Magdalena, Colombia; new synonymy]. times longer than phallobasic apodeme. Paramere: Schenkling's type-specimen does not differ sig- apical region papilliform in lateral view; lateral nificantly from the type of P. viridipennis. depression broad and well depressed, clearly visible DIAGNOSIS.—The gradual increase of flavotestace- in ventral view; dorsum convex and explanate ous color in the distal half of the antenna, and the medially; venter concave; mesoventral margin con- lack of deeply impressed punctations at the elytral cave; mesodorsal margin sinuous and denticulate. apical half easily distinguishes specimens of this Sinus: dorsal constricted at basal third, spheroid species from those of the other two species in the behind constriction, narrowly elliptical in front of viridipennis group. constriction. Phallus: marginal denticles of phallic DESCRIPTION.—Form: As in Figure 98. plate broad, shallow, and extended to phallic plicae Size: Length: males, average about 5.8 mm, range (9 specimens examined). 5.7-7.0 mm; females, average about 6.5 mm, range Female genitalia (Figures 100, 101): Dorsal lamina 5.9-7.6 mm. Width: males, average about 1.8 mm, bilobed, deeply incised; ventral lamina trilobed, range 1.6-2.1 mm; females, average about 2.2 mm, lateral lobes slightly shorter than medial lobe; range 1.8-2.4 mm. Ten males and 10 females meas- proctiger robust; proctigeral bacculi not fused; ured. ventral bacculi acuminate at posterior third; coxital Color: Cranium rufescent; antenna bicolorous, plates absent (1 specimen examined). basal half dark brown, distal half increases in pale- Internal Reproductive Organs: Male (Figures 107, ness; thorax rufescent except female pronotum (and 108); anterior accessory gland uniramous, tightly usually of males) with dark brown midapical mac- coiled; posterior accessory gland biramous, outer ula; mesoscutellum rufescent; femora flavotestace- branch one-third longer than inner branch; testis ous, usually infuscated dorsally; elytron variable composed of 12 follicles; basal third of ejaculatory (see "Variation"); abdominal venter rufescent. duct exceptionally bulbous. Female as in Figure 55 Head: Interocular depressions shallow; frontal (2 males and 1 female examined). umbo shallow; eyes moderately convex; HW/IOW, VARIATION.—Structural: The convexity of the average about 2.2, range 2.0-2.3 (10 males and 10 elytron seems to correlate geographically, the elytra females measured); antenna (Figure 97) serrate; being more convex in specimens from Panama than AL/PLS, 1.4; length/width ratio of each male anten- in those from Venezuela. nal article 2.4:1.4:1.8:1.8:1.5:1.3:1.2:1.1:1.0:0.93:1.5. Color: In some Panamanian specimens the mid- Pronotum: Only slightly transverse, rarely length apical macula of the pronotum extends posteriorly SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 96-103.—Perilypus insectus: 96, antenna. P. viridipennis: 97, antenna; 98, habitus; 99, elytral posthumeral region; 100, ovipositor, ventral view; 101, ovipositor, dorsal view; 102-103, elytra. (Scale = 1 mm) NUMBER 227 39

to form a narrow discal streak. Variation of elytral posited in BMNH, 1 in GEKI, and 1 in MNHP. here coloration suggests an east to west cline, but it may designated; type-locality: Volcan de Chiriquf, Provincia de also correlate with altitude as the few elevation Chiriquf, Panama]. records indicate. The lighter specimens were col- DIAGNOSIS.—Specimens of this species can be lected at low elevations in Panama, the darker easily confused with Panamanian specimens of P. ones at higher Andean elevations in Colombia and viridipennis, primarily because their elytral colora- Venezuela. Panamanian specimens are predomi- tion is very similar (compare Figures 103, 110). nantly rufescent and have a black to violaceous Anatomically, however, the two species are quite basal macula on the elytron (Figure 103). This distinct. Specimens of P. relucens are generally macula may be confined to the humeral angle or broader, their elytra less convex posteriorly and extend from the post humeral margin to the su- feebly depressed at middle third. The convexity of tured margin. In specimens from Colombia the their epipleural fold is almost obliterated; the elytron may be entirely blue, or blue in basal and fold is collapsed and appears grooved longitudi- apical thirds and rufescent in middle third (Figure nally. Also their elytral punctations are slightly 102). Venezuelan specimens have the elytron shiny smaller, extend to the suture, and terminate at blue or shiny blue-green. elytral posterior fourth. NATURAL HISTORY.—In May, on Cerro Campana, DESCRIPTION.—Form: As in Figure 110. Panama, I collected a female by beating woody Size; Length: males, average about 6.9 mm, range shrubs and lianas at an altitude of 460 meters. At 5.9-7.7 mm; females, average about 7.4 mm, range the same site and by the same collecting method, 6.1-8.0 mm. Width: males, average about 2.2 mm, H. Stockwell collected two male adults in January. range 2.0-2.4 mm; females, average about 2.5 mm, Other specimens were collected in Panama in range 2.2-2.8 mm. Ten males and 10 females February, March, May, June, July, and November, measured. and in Colombia in May. The specimens from Color: Cranium refescent; antenna refescent or Venezuela were collected in June by J. and B. brown; prothorax rufescent, with or without dark Bechyne\ one at 850 meters, the other at 1000 brown midapical macula; scutellum, meso- and meters. metasternum, legs, and abdomen rufescent or dark DISTRIBUTION (Figures 357, 358).—Specimens were brown; elytron either blue in basal fourth and collected in central Panama, northern Colombia, rufescent in apical three-fourths, or entirely blue and northern Venezuela. and with a violaceous or virescent tinge. LOCALITY RECORDS (Figures 357, 358).—I examined 21 adult Head: As in P. viridipennis except interocular specimens from Central and South America. PANAMA: width proportionally wider (HW/IOW, average Provincia de Code: El Valle (NMNH, 1 male and 1 female). about 1.9, range 1.9-2.1; 20 specimens measured); Provincia de Panama: Rio Diablo (GEKI, 1 male) ; Cerro Campana (GEKI, 1 male and 1 female; HPST, 1 male). eyes shallower; antenna (Figure 111) proportionally Cacahualito (CMPP, 1 male). Provincia de Colon: Portobelo longer (AL/PLS, 1.6); and length/width ratio of (GEKI, 1 male). Canal Zone: Summit (GEKI, 1 female); each male antennal article 2.0:1.2:1.8:1.6:1.4:1.4: 3 miles [4.8 km] west of Paraiso (GEKI, 1 female); Barro Col- 1.4:1.4:1.3:1.2:1.8. orado Island (WBIV, 1 male). COLOMBIA: Departamento de Pronotum: As in P. viridipennis except more Magdalena: Sevilla (MCZC, 1 male and 1 female); Aracataca (GEKI, 2 females; MCZC, 1 female); Rio Magdalena (DEIE, transverse (PL/PW, average about 0.87, range 0.77- 1 male; ZMAN, 1 female). Colombia (MSPI, 1 specimen, sex 0.98; 20 specimens measured), less convex, and not known). VENEZUELA: Estado de Carabobo: Las subapical depression more sinuous. Trincheras, Hacienda El Palmar (CMPP, 1 female); Cerro Elytron: As in P. viridipennis except less convex, Aquirre (GEKI, 1 male) . feebly depressed at middle third, punctations larger (Figure 109) and extended to posterior three- 6. Perilypus relucens (Gorham), new combination fourths, epipleural fold collapsed, and epipleural margin less convex at posterior third. FICURES 109-114,357 Front Tarsal Claws: As in P. viridipennis. Colyphus relucens Gorham, 1886:336 [lectotype: female, de- Male Genitalia (Figures 112-114): Strongly pig- posited in BMNH, and 4 paralectotypes, females, 2 de- mented. Combined length phallobase and phal- 40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 104-110.—Perilypus viridipennis: 104, tegmen, lateral view; 105, tegmen, ventral view; 106, phallus; 107, male internal reproductive organs; 108, posterior accessory glands. P. relucens: 109, elytral posthumeral region; 110, habitus. (Scale = 1 mm) NUMBER 227 41

Y----

113 114

FIGURES 111-117.—Perilypus relucens: 111, antenna; 112, tegmen, lateral view; 113, tegmen, ventral view; 114, phallus. P. levis: 115, tegmen, lateral view; 116, tegmen, ventral view; 117, phallus. (Scale = 1 mm) 42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY lobasic apodeme 7 times longer than paramere. MNHP, 2 males and 1 female); Las Lagunas (GEKI, 1 fe- Combined length of paramere and phallobase 3.6 male). "Panama" (ZMAN, 2 females). COLOMBIA: De- partamento de Valle del Cauca (GEKI, 1 male). times longer than phallobasic apodeme. Paramere: apex uncinate in ventral view; lateral depression REMARKS.—Gorham (1886:336), unaware of this broad, more basoventral than basolateral; dorsum species' sex linked color polymorphism, identified feebly convex and not explanate medially; venter the males as Colyphus signaticollis Spinola. He did, narrowly concave; mesoventral margin straight in however, note the anatomical similarity between the basal two-thirds, slightly arcuate in apical third; two sexes. In discussing the females he states, "Al- mesodorsal margin sinuous. Sinus: dorsal and ven- lied to the blue variety [= P. relucens males] of tral sagittate, ventral slightly narrower than dorsal. C. signaticollis figured in the earlier part of this Phallus: phallic plate narrowing from middle half; volume." marginal denticles restricted to apical half of phallic plate (10 specimens examined). 7. Perilyptis levis, new species Female Genitalia: As in P. viridipennis except proctiger broader and posterior margin more arcu- FIGURES 115-123,358 ate (1 specimen examined). Internal Reproductive Organs: Female, as in TYPE-LOCALITY—Santarem, Estado de Brasil. Figure 55 (1 specimen examined). TYPE-SPECIMENS.—The male holotype and allo- VARIATION.—Structural: No conspicuous struc- type are deposited in CMPP. Five paratypes: GEKI, tural variation was noted, except that the epi- 2 males and 1 female; CMPP, 1 male and 1 female. pleural fold is more collapsed in some specimens DIAGNOSIS.—Within the viridipennis group speci- than in others. mens of this species are easily recognized by their Color: Specimens from the type-locality are par- boldly serrate antenna (Figure 123). tially sex dimorphic in elytral and abdominal DESCRIPTION.—Form: As in Figure 120. coloration. In male specimens, the elytron is en- Size: Length: Males, average about 6.7 mm, range tirely blue and may have a purpurescent or viriscent 6.1-7.1 mm; females, average about 6.8 mm, range tinge, and the abdomen is dark brown. In females 6.5-7.8 mm. Width: males, average about 1.9 mm, the elytron is blue in the basal fourth and rufescent range 1.8-2.2 mm; females, average about 1.9 mm, in the apical three-fourths, and the abdomen is range 1.8-2.0 mm. Eight males and 9 females rufescent. In one female the elytron is entirely measured. blue and the abdomen predominantly brown. Color: Cranium usually flavotestaceous, some- In one of two other females from Panama (exact times dark brown or black; antenna dark brown or locality not known) the elytron is entirely blue and black, rarely flavous apically; pronotum ranging has a purpurescent tinge, and the abdomen is en- from predominantly flavotestaceous to predomi- tirely brown. In the second specimen the elytron is nantly black (see "Variation" and Figures 118-122); entirely rufescent except for a small blue macula on femur flavotestaceous, infuscated apically; tibia and the humeral angle. The abdomen is entirely rufes- tarsus dark brown; elytron violaceous, cupreus, or cent. blue-black, and with or without broad flavotesta- NATURAL HISTORY.—In May, in lakeside forest at ceous vitta on epipleural margin, rarely with faint Las Lagunas, Panama, I collected one female speci- discal vitta (epipleural-margin vitta extended to men by beating tree foliage. G. E. Champion col- apex or not); abdominal venter dark brown or lected adults on Volcan de Chiriquf at elevations black. from 610 to 1829 meters. Head: As in P. viridipennis except antenna DISTRIBUTION (Figure 358).—The known range (Figure 123) broader, more serrate (see "Varia- extends from northern Panama to western Colom- tion"), more setose, and proportionally longer bia. (AL/PLS, 1.6); and length/width ratio of each male antennal article 2.0:1.3:1.4:1.5:1.0:1.0:1.0:0.9: LOCALITY RECORDS (Figure 358) .—I examined 21 adult specimens from Central and South America. PANAMA: 0.8:0.8:1.3. Provincia de Chiriquf: Volcan de Chiriquf (BMNH, 4 males Pronotum: As in P. viridipennis except length and 3 females, lectotype; GEKI, 5 males and 2 females; equals width (7 specimens measured). NUMBER 227 43 Elytron: As in P. viridipennis except more slen- in specimens from Santarem, Brazil, and in females der (EL/EW, average about 4.7, range 4.4-5.2; 17 from Colombia and the Canal Zone (Figure 122). specimens measured), less convex, and punctation The pronotum (Figures 118-120) is predominantly extended to sutural margin and to elytral apical black in specimens from Amapa, Brazil, and in third (but see "Variation"). males from Venezuela and Colombia. Front Tarsal Claws: As in P. viridipennis. The male from Bogota, Colombia, has a short Male Genitalia (Figures 115-117): Feebly pig- and narrow flavotestaceous vitta on the elytral disc mented; tegmen spatulate. Combined length of and one on the epipleural margin. In specimens phallobase and phallobasic apodeme 4.7 times from Colombia, and in one of three females from longer than paramere. Combined length of para- the Canal Zone, the epipleural vitta is broader than mere and phallobase 2.5 times longer than phallo- the epipleural fold and extends to the elytral apex; basic apodeme. Paramere: very broad in lateral in other specimens the marginal vitta is as broad or view; dorsum boldly convex, explanate medially; narrower than the epipleural fold and terminates venter convex broad at base and tapered to apex; before the elytral apex. The antennae are uni- mesoventral margin sinuous; mesodorsal margin formly dark brown or black except in three females arcuate. Sinus: dorsal digitform but widened be- from the Canal Zone, in which they are increasingly tween paramere apices; ventral abconic, slightly flavotestaceous towards the apex. shorter than dorsal, and feebly constricted basally. NATURAL HISTORY.—From Barro Colorado Is- Phallus: phallic plates without marginal denticles land, Canal Zone, T. L. Erwin collected one adult (8 specimens examined). specimen in June. P. J. Darlington collected two Female Genitalia: As in P. viridipennis except additional adults from the same locality in May. lobes of ventral lamina equal in length (1 speci- Other specimens were collected from Colombia in men examined). September by M. de Ma than, from Brazil between Internal Reproductive Organs: Female, as in October and November by Thieme, and from Vene- Figure 55 (1 specimen examined). zuela in August by J. and B. Bechyn6, and between VARIATION.—Structural: Degree of antennal ser- September and October by E. A. Klages. ration, and apportionment of elytral punctations DISTRIBUTION (Figure 358).—This widely distrib- vary substantially. Antennal serration varies geo- uted species ranges from the Canal Zone to Brazil graphically, which is particularly evident among and is recorded from highlands of Colombia and male specimens; serration of the clavola is more Venezuela. prominent in eastern than in western South Ameri- ETYMOLOGY.—Latin, the adjective lews (smooth). can specimens. I refer to the smooth surface at the sutural and There is some indication that development and apical region of the elytron. distribution of elytral punctations is also correlated with geography; the. punctations are notably re- LOCALITY RECORDS (Figure S58).—I examined 17 adult duced or absent at elytral apical third in specimens specimens from Central and South America. CANAL ZONE: from Santarem (Brazil), Colombia, or Venezuela. Barro Colorado Island (GEK.I, 1 female; MCZC, 1 female; NMNH, 1 female). COLOMBIA: Departamento de Cundi Elytral punctation is conspicuously coarser and namarca; Bogota (GEKI, 1 male); Conanche (NMNH, 1 more densely distributed among the geographically female). Departamento de Boyaca; Muzco (GEKI, 1 male disjunct specimens from the Canal Zone and and 1 female). VENEZUELA: Distrito Bolivar: Suapure, Amapa, Brazil (the two specimens from Amapa are Rio Cauca (CUNY, 1 male). BRAZIL: Territorio de particularly coarsely punctated). I assign these Amapa: Sena Lombarda (USMV, 1 female); Rio Calcoene specimens to a deep-punctated morph. (GEKI, 1 male). Estado de Para: Santarem (GEKI, 2 males and 1 female; CMPP, 2 males, holotype, and 2 fe- Color: Color polymorphism appears to correlate males) . geographically and possibly altitudinally. Specimens from lower elevations (Canal Zone and Santarem, REMARKS.—The magnitude of infraspecific varia- in Brazil) have a lighter pronotum than those col- tion involving elytral punctation is quite excep- lected from the more mountainous regions of Co- tional. Perhaps the specimens from the Canal Zone lombia and Venezuela. and Amapa, Brazil, are not conspecific with the The pronotum is predominantly flavotestaceous other specimens examined, but the genitalic char- SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 44

FICURES 118-124.—Perilypus levis: 118-119, 121-122, pronotum; 120, habitus; 123, antenna. P. limbatus: 124, antenna. (Scale = 1 mm) NUMBER 227 45 acteristics in the male from Amapa are identical decoris, have the anterior margin of antennal article with males whose elytral punctation is typical. I 9 to 11 flavotestaceous. Perilypus limbatus differs assume that within the viridipennis group homo- from P. apocopatus by having the femora predomi- geneity of genitalic characteristics indicates con- nantly flavotestaceous, and from P. decoris by the specificity, as they do in other species groups of the narrower epipleural fold and absence of the elytral genus; perhaps the observed differences are early discal vitta. indications of peripheral isolation. DESCRIPTION.—Form: As in Figure 125. Although I did not study male specimens from Size: Length: males, average about 6.9 mm, range the Canal Zone, I tentatively consider the three 6.3-7.6 mm; females, average about 8.2 mm, range female specimens from that region as conspecific. I 7.2-9.2 mm. Width: males, average about 1.9 mm, labeled them "Perilypus levis Ekis?" range 1.8-2.0 mm; females, average about 2.4 mm, range 2.1-2.8 mm. Ten males and 10 females The limbatus Group measured. Color: Clypeus, frons (Figure 57), and venter of The following combination of characteristics dis- cranium flavotestaceous; epicranium and gena shiny tinguish members of the limbatus group from other black; antenna predominantly black, anterior mar- Perilypus: body form rectangulate; antenna moder- gin of articles 9 to 11 flavotestaceous, pronotum ately serrate (flagellar articles subfiliform in males); variable (see "Variation"); legs bicolorous; femur elytron with convex epipleural fold, deeply im- flavotestaceous, with dorsoapical infuscation dimin- pressed and widely distributed punctations, punc- ishing from front to hind femur; tibia and tarsus tate and interpunctate surfaces smooth and shiny, black to brown; prosternum flavotestaceous, meso- and with a gradual apical slope. and metasternum black to brown; elytron variable The composite distribution of the 10 species of (see "Variation"); abdomen black. this group extends from Chiapas, Mexico, to San Head: Interocular depressions shallow; frontal Martin, Peru; the range north of Panama is at- umbo indistinct; eyes boldly convex (HW/IOW, tributed only to locality records of P. prolixipenis. average about 2.2, range 2.1-2.4; 20 specimens measured); antenna (Figure 124) moderately serrate; 8. Perilypus limbatus (Gorham), new combination AL/PLS, 1.7; length/width ratio of each male antennal article 2.5:1.4-1.7:1.7:2.0:1.5:1.4:1.3:1.1:1.0 FIGURES 124-132, 136, 137, 362 1.9. Colyphus limbatus Gorham, 1878:161 [lectotypc: male, here Pronotum: Only slightly transverse; PL/PW, designated, deposited in MNHP; type-locality: Caracas, average about 0.93, range 0.89-0.96 (20 specimens Venezuela]; 1886:336. measured); subapical depression deeply impressed Derestenus limbatus var. latesuturalis Pic, 1941:11 [lectotype: and strongly sinuous; pronotal arch impressed with female, here designated, deposited in MNHP; type-locality: few shallow punctations; side margin of pronotum Bolivia; new synonymy]. proper moderately arcuate. Derestenus limbatus var. notaticollis Pic, 1941:11 [lectotype: female, here designated, deposited in MNHP; type-locality: Mesoscutellum: Subquadrate. Venezuela; new synonymy]. Elytron: Epipleural margin straight in basal Derestenus limbatus var. suturalis Pic, 1941:11 [lectotype: three-fourths, arcuate in apical fourth; apical slope male, here designated, deposited in MNHP; type-locality: gradual; disc impressed with large punctations at Venezuela; new synonymy]. basal two-thirds, punctations are smaller at apical Derestenus columbicus var. reynoldsi Wolcott, 1927b: 106 [holotype not studied; see "Remarks," one paratype in third; punctate and interpunctate surfaces smooth FMNH; type-locality: El Valle, Venezuela; new synonymy]. and shiny; epipleural fold convex; EL/EW, average Derestenus latefasciatus Pic, 1941:10 [lectotype: female, here about 5.1 range 4.6-5.7 (20 specimens measured). designated, deposited in MNHP; type-locality: Bolivia; new Front Tarsal Claws: Male, strongly asymmetrical. synonymy]. Female, feebly asymmetrical. The names in synonymy represent intrapopula- Male Genitalia (Figures 128-130): Strongly pig- tion color variants. mented. Combined length of phallobase and phal- DIAGNOSIS.—Within the limbatus group only lobasic apodeme 4 times longer than paramere. specimens of P. limbatus, P. apocopatus, and P. Combined length of paramere and phallobase 4.8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 46

FIGURES 125-130.—Perilypus limbatus: 125, habitus; 126, pronotum; 127, elytra; 128, aedeagus, ventral view; 129, phallus, ventral view, X 130 (arrow indicates marginal denticles); 130, phallus, lateral view, X 130 (arrow indicates phallic plicae. (Scale = 1 mm) NUMBER 227 47 times longer than phallobasic apodeme. Paramere: the epipleural margin and one on the sutural Apex obtuse; lateral depression present; dorsum margin. In some specimens of the bimaculate and convex, explanate medially; venter concave; meso- bivittate phena the pronotum has a fuscous testa- ventral margin straight in basal half, faintly sinuous ceous macula anteriad to the fovea. In one speci- in distal half; mesodorsal margin sinuous. Sinus: men of the bivittate phenon, the marginal vitta dorsal lanceolate, constricted subapically; ventral extends only to the elytral basal half. In specimens obconic. Phallus: apex papilliform; phallic plates of the punctate phenon, the pronotum is fulvous to contiguous dorsally; marginal denticles stout and fulvescent and has a black midapical macula and extended beyond limit of phallic plicae (20 speci- black collar (Figure 125); the pronotal macula mens examined). varies in width. Members of the fasciate phenon Female Genitalia (Figures 131, 132): Dorsal have a punctate pronotum and a broad flavous lamina bilobed; ventral lamina trilobed, lateral discal fascia on the midelytron (Figure 125). The lobes shorter than medial lobe; proctiger reduced; entire spectrum of color polymorphism described proctigeral bacculi not fused; ventral bacculus above is present in one population sample from acuminate at posterior fourth; coxital plates promi- Rancho Grande, Venezuela. nent; coxital stylus bulbous distally (2 specimens Seven specimens from El Lim6n, Venezuela, are examined). more melanistic than the other specimens exam- Internal Reproductive Organs: Male (Figures ined. They exhibit the following characteristics: 136, 137), anterior accessory gland uniramous, posterior region of clypeus black, femur black tightly coiled; posterior accessory gland biramous, distally, and elytral vitta absent or indistinct. outer branch about one-fourth longer than inner NATURAL HISTORY.—In May, in the vicinity of branch; testis composed of 12 follicles. Female, as the Rancho Grande Biological Station in Aragua, in Figure 55; ovary composed of 12 ovarioles (5 Venezuela, I collected 53 adults, 2 larvae, and 1 males and 3 females examined). pupa. The macrohabitat of the site can be described VARIATION.—Structural: The degree of concavity briefly as a predominantly deciduous, subtropical of the elytral epipleural margin is notably variable. montane forest. The topographical location of this Females from El Valle, Venezuela, have the epi- Andean region, generally considered a cloud forest, pleural margin conspicuously more concave than affords some protection from the nubilous condi- do females from other localities; El Valle females tions generated by the Caribbean Ocean. Conse- are also more robust in body form. quently the collection site (at about 1100 meters) Color: The available specimens, although gen- is considerably drier than adjacent mountainous erally homogeneous in external structure, are areas, particularly those at higher elevations. I did chromatically polymorphic. Pronotal and elytral not find P. limbatus specimens at 1200 meters pigmentation segregate specimens into four color where clouds maintained moisture level at con- phena: the bimaculate phenon (with reference to stant saturation. pronotal lateroapical maculae), the bivittate Woody lianas and low canopy verdure are two phenon (with reference to bivittate elytron), the of the site's most characteristic types of vegetation. punctate phenon (with reference to pronotal and Adults and one larva were collected by beating apical macula), and the fasciate phenon (with ref- vegetation assemblages rich in lianas of about 7-13 erence to fasciate elytron). Both sexes are repre- cm in diameter. I found a second larva and one sented by the first two phena, the last two only by pupa after examining numerous sections of dead females. lianas that housed the immatures in a shallow To the bimaculate phenon I assign individuals cavity beneath thin bark. On the same kind of that have a flavotestaceous lateroapical macula on liana I observed a female oviposit into a crevice in each anterior angle of the pronotum (as in Figure which I subsequently found one egg. Although I 157) and whose elytron is vittate at the epipleural failed to rear the aforementioned immatures to margin. In two female specimens of this phenon imago stage I am confident that they are members the pronotal disk is flavotestaceous. Specimens of of P. limbatus. The pupa (Figures 380, 381) albeit the bivittate phenon have a bimaculate pronotum, not mature, clearly exhibits typical characteristics and each elytron (Figure 127) has a yellow vitta on of adult Perilypus. The color markings of the 48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY pupal abdomen are identical to that found in the Wolcott's description leaves little doubt that the larvae; and I do not know of any other species of synonymy is correct; the holotype of this variety Perilypus collected from Rancho Grande vicinity. is assumed lost (Henry H. Dybas, of the Field At Rancho Grande I also collected adults of this Museum of Natural History, pers. comm.). species by black light; however, the species apparently is only mildly phototropic since only three 9. Perilypus columbicus (Spinola), new combination specimens were collected. The altitudinal range of P. limbatus, as indi- FIGURES 133-135, 138, 139, 362 cated by label data, is from 900 to 1500 meters. Thanasimus columbicus Spinola, 1844a: 195 [lectotype: fe- DISTRIBUTION (Figure 362).—I studied specimens male, and one paralectotype, male, subsequent designa- from northern Venezuela and Bolivia. The seem- tion by Elds, 1975:27, lectotype deposited in MNHP, para- ingly disjunct distribution is probably a conse- lectotype in MSPI; type-locality: Bogota, Colombia].— quence of coincidental collecting. The affinity of Chevrolat, 1876:46,—Schenkling, 1906:263,—Ekis, 1975:27. Clerus colotnbiae Spinola, 1844b:135 [lectotype: female, here this species for subtropical montane forests sug- designated, deposited in MNHP; type-locality: Colombia; gests that they occur along the Atlantic slopes of new synonymy].—Gemminger and Harold, 1869:1735.— the Andes, which would make a transamazonian dis- Chevrolat, 1876:46. tribution unlikely. Spinola's (1844b: 135) lectotype represents a LOCALITY RECORDS (Figure 362).—I examined 122 adults, color phenon of P. columbicus. 2 larvae, and 1 pupa from South America. VENEZUELA: Distrito Federal: Caracas (ZMAN, 1 male and 1 female; DIAGNOSIS.—In addition to the characteristics FMNH, 1 male and 1 female; GEKI, 2 males; MNHP, 1 given in the key, specimens of this species are dis- male, lectotype; NMNH, 1 male); Quebrada Avila (CBAZ, tinguished from other members of the limbatus 1 male); Rio Gurimare (GEKI, 1 female); El Lim6n, Cordil- group by the proportionally shorter antenna lera Del Litoral (CBAZ, 2 males and 1 female; GEKI, 2 (AL/PLS, 1.5) and by the length/width ratio of males and 2 females); El Valle (FMNH, 2 females; GEKI, 1 female). Estado de Carabobo: Cerro de Caffe (GEKI, 1 pronotum (0.99). Also, the lateral explanation of female); Santa Clara (UCMV, 1 male); Trincheras (GEKI, 1 the phallobase and the basal region of the lateral female, UCMV, 1 female). Estado de Aragua: Maracay, lobes (Figure 134) is a modification of the male Rancho Grande (AMNH, 2 males and 1 female; ZMAN, 3 genitalia unique in Perilypus. males; BMNH, 3 males and 1 female; CBAZ, 3 males and 1 female; FMNH, 4 males; GEKI, 18 males, 19 females, 1 DESCRIPTION.—Form: As in P. limbatus (Figure larva and 1 pupa; CMPP, 2 males; UCMV, 2 males and 2 125). females; MNHP, 2 males and 1 female; MCZC, 2 males and Size: Length: male, average about 7.6 mm, range 1 female; MZSP, 2 males; NMNH, 13 males, 4 females and 7.4-7.9 mm; female, average about 7.7 mm, range 1 larva). "Venezuela" (FMNH, 1 female, MNHP, 5 fe- 7.1-8.3 mm. Width: male, average about 2.2 mm, males) . BOLIVIA: Departamento de Cochabamba: Cocha- bamba (GEKI, 1 female). "Bolivia" (GEKI, 1 male; MNHP, range 2.1-2.4 mm; female, average about 2.4 mm, range 2.0-2.6 mm. Seven specimens measured. 3 males and 1 female). Color: Clypeus, gula, prosternum, and basal half REMARKS.—The new synonymies are based on of femur flavotestaceous; labrum, frons, cranium, comparisons of the appropriate type-specimens distal half of femur, tibia, tarsus, meso- and meta- and on available biological data; I did not study sternum, and abdomen black; pronotum and ely- the type of Wolcott's variety. The color variants, tron variable (see "Variation"). herein assigned to phena, were given varietal or Head: As in P. limbatus. species status by previous authors. The varietal Pronotum: As in P. limbatus except feebly trans- names of Pic (1941), latesuturalis, notaticollis, verse (PL/PW, average about 0.99, range 0.95-1.0 suturalis, and that of Wolcott (1927b), reynoldsi, (7 specimens measured). correspond to the bimaculate, bivittate, punctate, Elytron: As in P. limbatus except EL/EW, aver- and fasciate phenon, respectively. Further, Dereste- age about 4.5, range 4.3-4.9 (7 specimens meas- nus latefasciatus Pic, corresponds to my fasciate ured). phenon and to Wolcott's reynoldsi variety. I did not Front Tarsal Claws: As in P. limbatus. study the holotype specimen of D. columbicus var. Male Genitalia (Figures 133-135): Strongly pig- reynoldsi Wolcott although a para type and mented. Combined length of phallobase and phallo- NUMBER 227 49

FIGURES 131-141.—Perilypus limbatus: 131, ovipositor, ventral view; 132, ovipositor, dorsal view. 136, male internal reproductive organs; 137, posterior accessory glands. P. columbicus: 133, tegraen, lateral view; 134, tegmen, ventral view; 135, phallus; 138, ovipositor, ventral view; 139, ovipositor, dorsal view. P. iris: 140, ovipositor, ventral view; 141, ovipositor, dorsal view. (Scale = 1 mm) 50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY basic apodeme 4 times longer than paramere. 10. Perilypus apocopatus, new species Combined length of paramere and phallobase 3.1 times longer than phallobasic apodeme. Paramere: Figures 143-148, 362 explanate basally; apical region feebly arcuate in TYPE-LOCALITY.—Ocana, Departamento de None lateral view; lateral depression small but well im- de Santander, Colombia. pressed; dorsum convex, explanate medially; venter TYPE-SPECIMENS.—The holotype male is depos- concave; mesoventral margin concave; mesodorsal ited in MNHP, the allotype in GEKI. margin sinuous. Sinus: dorsal narrow and almost DIAGNOSIS.—Distal half of hind femur predomi- twice as long as ventral, ventral elliptical. Phallo- nantly black; inner margins and venter of antennal base: explanate in apical half. Phallus: acuminate; articles 9 to 11 testaceous. Elytral coloration of the marginal denticles as in P. limbatus (3 specimens P. apocopatus female specimen is similar to that of examined). fasciate females of P. limbatus and P. columbicus, Female Genitalia (Figures 138, 139): As in P. except that the elytral fascia of P. apocopatus does limbatus except lateral lobes of ventral lamina as not reach the sutural margin. long as medial lobe and proctiger more prominent. DESCRIPTION.—Form: As in P. limbatus (Figure VARIATION.—Structural: The beetles examined 125). are structurally homogeneous externally. Size: Length: male, 7.4 mm; female, 7.0 mm. Color: The male pronotum and elytron are Width: male, 2.2 mm; female, 2.2 mm. One male black, except the former is fuscous lateroapically, and one female measured. the latter feebly vittate on the epipleural margin. Color: As in P. limbatus except posterior half of Two color patterns of the female elytron distin- frons black; antennal articles 9 to 11 predominantly guish a vittate phenon and a fasciate phenon. flavotestaceous; and hind femur black in distal half. Pronotal and elytral coloration varies with Vittate females have the elytron black; a faint sex (see "Variation"). vitta on the epipleural margin may be present. In Head: As in P. limbatus. fasciate females, the elytron has a broad rufescent Pronotum: As in P. limbatus. fascia at the midelytron which extends to the Elytron: As in P. limbatus except EL/EW, aver- sutural margin, and a broad rufescent macula is age about 4.5, range 4.4-4.6 (2 specimens present on the elytral apex. measured). NATURAL HISTORY.—Unknown, except that two Front Tarsal Claws: As in P. limbatus. specimens were collected from Bogota, a locality Male Genitalia (Figures 146-148): Strongly pig- at about 4000 meters on the Cordillera Oriental mented. Combined length of phallobase and phallo- of the Colombian Andes. basic apodeme 3 times longer than paramere. DISTRIBUTION (Figure 362).—Known only from Combined length of paramere and phallobase 4.0 Colombia. times longer than phallobasic apodeme. Paramere: apical region arcuate and acuminate in lateral LOCALITY RECORDS (Figure 362).—I examined 6 adult specimens from South America. COLOMBIA: Departamento de view; lateral depression feebly developed; dorsum Cundinamarka: Bogota (MNHP, 2 females, lectotype; MSPI, convex, broadly explanate medially; venter con- 1 male). "Colombia" (GEK.I, 2 males and 1 female). cave; mesoventral and mesodorsal margin concave. Sinus: dorsal narrow, digitiform; ventral excep- REMARKS.—I (Ekis, 1975:25) erroneously con- tionally long and elliptical. Phallus: apical region cluded that the specimen in MSPI was the holotype. acutely narrowed; phallic plates narrowed in With regard to Clerus colombiae Spinola, Chevro- apical half; marginal denticles as in P. limbatus. lat (1876:46) poses the question "C'est un Female Genitalia (Figures 143, 144): Dorsal Colyphus?" He was quite correct in doubting the lamina broadly incised; ventral lamina reduced, placement of this species under Clerus (now Eno- lateral lobes longer than very shallow medial lobe; clerus). The width of the distal articles of colom- oblique bacculus incorporated into large anterior biae antennae is exaggerated in the habitus illus- coxital plate; proctiger short; proctigeral and ven- tration published by Spinola (1844b, pi. 6; fig. 6). tral bacculi as in P. limbatus. NUMBER 227 51 VARIATION.—Structural: The specimens examined apically (Figure 145); and the nonflavotestaceous are structurally homogeneous externally. portions of the elytron are more purpurescent than Color: The two specimens studied differ in pro- black. notal and elytral coloration. In the male pronotum DISTRIBUTION (Figure 362).—Known only from is predominantly black and fuscous lateroapically; Ocana, located on the northern latitude of the the elytron is black, with a purpurescent tinge, and Colombian Cordillera Oriental. is broadly flavotestaceous apically. In the female ETYMOLOGY.—A Latin derived adjective from the pronotum is predominantly flavotestaceous and the noun apocope (a cutting off) + the suffix has a black midapical macula and black pronotal -atus (cut off). The name refers to the abbreviated collar; the elytron has a broad flavotestaceous fascia length of the dorsal sinus of the male genitalia. at the middle and an epipleural vitta that expands

142

FIGURES 142-145.—Perilypus iris: 142, habitus. P. apocopatus: 143, ovipositor, ventral view; 144, ovipositor, dorsal view; 145, elytra. (Scale = 1 mm) SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 52

LOCALITY RECORDS (Figure 362).—I examined 2 adult paratypes are deposited in repositories as noted specimens from South America. COLOMBIA: Departamento under "Locality Records." de Norte de Santander: Ocaiia (GEKI, 1 female; MNHP, 1 DIAGNOSIS.—Distinguishable from the very simi- male, holotype). lar specimens of P. limbatus by the following combination of characteristics: front tarsal claws sym- 11. Perilypus iris, new species metrical, antennal articles 9 to 11 not testaceous, lateroapical maculae of pronotum (when present) FIGURES 140-142, 362 extended posteriorly beyond subapical depression. TYPE-LOCALITY.—Tocota, Departamento de Valle DESCRIPTION.—Form: As in P. limbatus (Figure del Cauca, Colombia. 125). TYPE-SPECIMENS.—The holotype female is depos- Size: Length: males, average about 7.2 mm, range ited in ZMAN. 6.1-7.9 mm; females, average about 7.5 mm, range DIAGNOSIS.—Within the limbatus group speci- 6.4-8.4 mm. Width: males, average about 2.0 mm, mens belong to this species if their femora are range 1.8-2.1 mm; females, average about 2.2 mm, entirely black. range 2.0-2.3 mm. Twelve males and 5 females DESCRIPTION.—Form: As in Figure 142; forebody measured. narrow; elytra convex. Color: Head, thoracic sterna, legs, and abdomen Size: Female: length, 8.3 mm; width, 2.5 mm. as in P. limbatus, or postocular region of cranium Color: Head, meso- and metasternum, legs, and flavotestaceous; antennal articles 9 to 11 entirely abdominal venter black; prothorax flavotestaceous, black; pronotum predominantly black, lateroapical except pronotal arch with transverse black macula; flavotestaceous maculae (when present) extended elytron with broad sharply outlined fascia at mid- beyond subapical depression, and discal vitta per- dle, and at apex. current or not (Figure 149); and vitta on Head: As in P. limbatus. epipleural margin broader. Pronotum: As in P. limbatus except feebly trans- Head: As in P. limbatus. verse (PL/PW, 0.99). Pronotum: As in P. limbatus. Elytron: As in P. limbatus except more convex Elytron: As in P. limbatus. and with more depth. Front Tarsal Claws: Symmetrical in both sexes. Front Tarsal Claws: Female, feebly asymmetrical. Male Genitalia (Figures 154, 155): Strongly pig- Female Genitalia (Figures 140, 141): Ventral mented. Combined length of phallobase and phallo- lamina with lateral lobes shorter than medial lobe; basic apodeme 5.5 times longer than paramere. oblique bacculus incorporated into small anterior Combined length of paramere and phallobase 4.5 coxital plate; coxite well sclerotized throughout; times longer than phallobasic apodeme. Paramere: dorsal lamina and proctigeral and ventral bacculi acuminate; dorsum plane, explanate medially; as in P. limbatus. venter concave; mesoventral margin arcuate; meso- ETYMOLOGY.—Latin, the noun iris (rainbow); dorsal margin feebly convex and denticulate. Sinus: with reference to the dorsal coloration of this dorsal narrowly obconic; ventral broadly elliptical. beetle. Phallus: apex lobifonn; phallic plates widely LOCALITY RECORDS (Figure 362).—I examined 1 adult speci- separated dorsally with marginal scales near apex men from South America. COLOMBIA: Departamento de (note arrow in Figures 154, 155) (12 specimens Valle del Cauca: Tocota (ZMAN, 1 female, holotype). examined). Female Genitalia (Figures 150, 151): As in P. 12. Perilypus acus, new species limbatus; except coxites more robust; dorsal lamina deltoid, with short broad incision; lateral lobes of FICURES 149-151,154, 155, 362 ventral lamina very short; and proctiger more TYPE-LOCALITY.—Callanga, Provincia de Cusco, prominent (2 specimens examined). Peru. VARIATION.—Structural: The specimens examined TYPE-SPECIMENS.—The holotype male is depos- are structurally homogeneous externally. ited in MNHB, the allotype in FMNH. Fourteen Color: Pronotal coloration is sex dimorphic. The NUMBER 227 53

FIGURES 146-155.—Perilypus apocopatus: 146, tegmen, lateral view; 147, tegmen, ventral view; 148, phallus. P. acus: 149, pronotum; 150, ovipositor, ventral view; 151, ovipositor, dorsal view; 154, phallus, lateral views, X 130 (arrow indicates phallic scales); 155, phallus, lateral view, X 1000 (upper arrow indicates phallic papillae; lower arrow indicates phallic scale). P. buga: 152, ovipositor, ventral view; 153, ovipositor, dorsal view. (Scale = 1 mm) SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 54 female pronotum is predominantly flavotestaceous more. Color of pronotum and elytron variable (see and has a black midapical macula and black "Variation"). pronotal collar. However, in three female speci- Head: As in P. limbatus except frontal umbo mens the black basal coloration of the pronotum more prominent; antenna proportionally longer extends anteriorly (Figure 149). In males, the (AL/PL, 1.9); and length/width ratio of each male pronotum is always predominantly black and the antennal article, 2.5:1.1:1.7:1.6:1.7:1.7:1.5:1.6:1.6: vitta on the elytral epipleural margin is narrower 1.3:2.0. than in female specimens. Pronotum: As in P. limbatus except PL/PW, NATURAL HISTORY.—Adults were collected in average about 0.91, range 0.88-0.95 (5 specimens Peru in May and August. measured). DISTRIBUTION (Figure 362).—The known distri- Elytron: As in P. limbatus except elytral costae bution extends from Tarapoto to Marcapata, Peru. prominent; vertical setae pale; and EL/EW, aver- ETYMOLOGY.—Latin, the noun acus (husk). I age about 5.0, range 4.7 to 5.4 (5 specimens refer to the scales on the apical region of the measured). phallic plates. Front Tarsal Claws: As in P. limbatus. Male Genitalia (Figures 158-160): Strongly pig- LOCALITY RECORDS (Figure 362).—I examined 16 adult spec- mented. Combined length of phallobase and phallo- imens from South America. PERU: Provincia de Cusco: basic apodeme 6 times longer than paramere. Callanga (FMNH, 1 male and 1 female; GEKI, 2 males and Combined length of paramere and phallobase 3.4 1 female; MNHB, 1 male, holotype); Marcapata (ZMAN, 1 male). Provincia de San Martin: Tarapoto (GEKI, 1 male times longer than phallobasic apodeme. Paramere: and 1 female; MNHP, 2 males and 1 female). "Peru" apex uncinate; lateral depression broad, conspicu- (BMNH, 1 male; GEKI, 2 males; MNHP, 1 female; ZMAN, ously long and most prominent near parameral 1 male). base; dorsum plane, only slightly explanate medially; venter concave; mesoventral margin con- 13. Perilypus buga, new species cave; mesodorsal margin straight. Sinus: dorsal elliptical; ventral abovate; both sinuses equal in FICURES 152,153, 156-162, 362 length. Phallus: marginal denticles small, barely extended to phallic plicae (1 specimen examined). TYPE-LOCALITY.—Buga, Valle del Cauca, De- Female Genitalia (Figures 152, 153): As in P. partamento de Valle del Cauca, Colombia. limbatus except coxites shorter, coxital stylus not TYPE-SPECIMENS.—The holotype male and allo- bulbous distally; proctiger more prominent; and type are deposited in NMNH. These and three dorsal lamina with very short medial lobe (2 speci- paratypes were collected by me in 1973. Three para- mens examined). types: GEKI, 3 females. Internal Reproductive Organs: Male (Figures DIAGNOSIS.—In addition to characteristics given 161, 162); anterior accessory gland uniramous, in the key, specimens of this species are distin- tightly coiled; posterior accessory gland biramous, guished from specimens of other species in the outer branch only slightly longer than inner limbatus group by the longer antenna (AL/PL, 1.9), branch; testis composed of 12 follicles; ejaculatory more conspicuous elytral costae, and by the pre- duct conspicuously bulbous in proximal half. Fe- ponderance of pale erect setae on the elytral male as in Figure 55; ovary composed of 12 ovari- surface. oles (1 male and 4 females examined). DESCRIPTION.—Form: As in Figure 156. VARIATION.—Structural: Conspicuous structural Size: Length: male, 5.9 mm; female, average variation appears to be restricted to convexity of about 7.6 mm, range 7.3-7.8 mm. Width: male the epipleural fold, which is most developed in the 1.5 mm; female, average about 2.2 mm, range 2.0- vittate female. 2.3 mm. One male and 4 females measured. Color: Sexual dimorphism with respect to pro- Color: As in P. limbatus except venter of anten- notal and elytral color is present. Color variation nal articles 9 to 11 black, venter of cranium and groups females into a punctate or bivittate phenon. posterior region of frons totally or predominantly These phena are generally similar to their P. lim- black (Figure 58), and femur black in apical half or batus counterparts, but in this species the pronotum NUMBER 227 55

156

FIGURES 156-163.—Perilypus buga: 156, habitus; 157, pronotum; 158, tegmen, lateral view; 159, tegmen, ventral view; 160, phallus; 161, male internal reproductive organs; 162, posterior accessory gland. P. latilira: 163, elytron (arrow indicates epipleural fold). (Scale = 1 mm) SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 56 of punctate specimens (Figure 156) is black along P. latilira are distinguished from specimens of the lower sides and at the base anteriad to the P. prolixipenis by male genitalic characteristics pronotal collar. The posthumeral elytral vitta is (compare Figures 168, 173) and probably by geo- faintly visible or absent in punctate females. The graphic distribution. elytron of the bivittate phenon (Figure 156) has a DESCRIPTION.—Form (Figure 165): A broad and broad discal vitta that does not join the marginal deep-bodied species. vitta at the elytral apex. In the male, color of the Size: Length: males, average about 7.6 mm, range pronotum is similar to Figure 157; the elytron is 6.4-8.9 mm; females, average about 8.6 mm, range black and avittate. 6.5-9.8 mm. Width: males, average about 2.0 mm; NATURAL HISTORY.—In May, in Buga, Colombia, range 1.9-2.5 mm; females, average about 2.6 mm, at an altitude of 1700 meters, 1 collected 5 speci- range 1.9-3.0 mm. Ten males and 3 females mens of this species by beating tree branches inter- measured. twined with small woody lianas. Color: Frons and venter of cranium flavotesta- DISTRIBUTION (Figure 362).—Specimens were col- ceous; gena black; epicranium with black subquad- lected from Buga, a location between Cordillera rate macula; antenna black; pronotal disc usually Occidental and Cordillera Central of the Colom- with black vitta that extends to hind angles of bian Andes. pronotum proper; lower sides of pronotum black; ETYMOLOGY.—"Buga" is the name of the locality femur flavotestaceous but black or infuscated dor- from which these beetles were collected. The soapically; tibia and tarsus dark brown; prosternum epithet is a noun in apposition. flavotestaceous; meso- and metasternum black; LOCALITY RECORDS (Figure 362).—I examined 5 adult elytron piceous, with purpurescent tinge and with specimens from South America. COLOMBIA: Departa- broad flavotestaceous marginal vitta that expands mento de Valle del Cauca: Valle del Cauca, Buga (GEKI, 3 apically; elytron in male with or without short females; NMNH, 1 male, holotype, and 1 female). and narrow flavotestaceous discal vitta, in female REMARKS.—The avittate females are superficially vitta (Figure 166) is broad and long and not con- very similar to the darker P. limbatus specimens fluent with vitta on epipleural margin; abdominal from El Limon, Venezuela. However, in P. buga venter in male black, in female flavotestaceous ex- specimens the elytral vertical setae are pale, not cept visible sterna 5 and 6 black. black as in El Limon specimens. The black colora- Head: As in P. limbatus except eyes less convex tion on the pronotal lower sides and at the base (HW/IOW, average about 2.2, range 1.8-2.4, 13 beyond the pronotal collar (Figure 156) further specimens measured) and antenna (Figure 164) distinguish P. buga specimens from the more mela- proportionally shorter (AL/PL, 1.4). nistic specimens of P. limbatus. Pronotum: As in P. limbatus except more convex, subapical depression more sinuous, and pronotal arch shallower and more prolonged medially. 14. Perilypus latilira, new species Elytron: As in P. limbatus, except broader and

FIGURES 164-171, 361 deeper (EL/EW, average about 4.9 range 4.5-5.4; 13 specimens measured) and epipleural fold (Fig- TYPE-LOCALITY.—Bugaba, Provincia de Chiriqui, ure 163) wider. Panama. Front Tarsal Claws: As in P. limbatus. TYPE-SPECIMENS.—The holotype male and the Male Genitalia (Figures 167-169): Strongly pig- allotype are deposited in BMNH. These and 14 mented. Combined length of phallobase and phallo- paratypes were collected from the type-locality by basic apodeme 6 times longer than paramere. G. C. Champion. The 14 paratypes are deposited Combined length of paramere and phallobase 3.4 in repositories as noted under "Locality Records." times longer than phallobasic apodeme. Paramere; DIAGNOSIS.—The combination of robust body, feebly arcuate in lateral view; dorsum narrow and epipleural fold wider than pedicel, and epicranium feebly convex; venter conspicously broader than with black subquadrate macula separate specimens dorsum, feebly concave; mesoventral and meso- of this species from those of other species in the dorsal margin concave. Sinus: dorsal and ventral limbatus group except P. prolixipenis. Specimens of elliptical, former broader. Phallus: acuminate; NUMBER 227 57

FIGURES 164-171.—Perilypus latilira: 164, antenna; 165, habitus; 166, elytra; 167, tegmen, lateral view; 168, tegmen, ventral view; 169, phallus; 170, ovipositor, ventral view; 171, ovipositor, dorsal view. (Scales = 1 mm) 58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY marginal denticles broad, shallow, and restricted TYPE-SPECIMENS.—The holotype male, deposited in middle region of phallic plate (13 specimens in FMNH, was collected by A. Hayne in 1922. Four examined). paratypes: deposited in repositories as noted under Female Genitalia (Figures 170, 171): Dorsal "Locality Records." lamina deeply incised and acuminate; lateral lobes DIAGNOSIS.—Within the limbatus group speci- of ventral lamina only slightly shorter than medial mens of this species are reliably separated from the lobe; proctigeral bacculi fused as in Figure 171; externally similar specimens of P. latilira, by male inner margin of coxite well sclerotized; coxital genital characteristics (compare Figures 168 and plates not visible (1 specimen examined). 174). VARIATION.—Structural: The epipleural fold is DESCRIPTION.—Form: As in P. latilira (Figure conspicuously broader and more convex in females 165). (Figure 163) than in males. The terminal article of Size: Length: males, average about 8.0 mm, the male antenna varies in length and in extent of range 7.4-8.9 mm; female, 9.8 mm. Width: males, acumination. average about 2.2 mm, range 2.0-2.5 mm; female, Color: Color variation is evident in both males 3.0 mm. Four males and 1 female measured. and females. One male specimen from Bugaba, Color: As in P. latilira. Panama, is particularly atypical; its cranium, an- Head: As in P. latilira. tenna, prothorax, and mesosternum, and legs are Pronotum: As in P. latilira. entirely flavotestaceous. Definition of the elytral Elytron: As in P. latilira. discal vitta is correlated with sex. Two of three Front Tarsal Claws: As in P. latilira. females examined have broad, clearly defined vittae Male Genitalia (Figures 172-174): As in P. lation the elytral disc (Figure 166). The elytral discs lira except combined length of phallobase and of the other female and some males show a faint phallobasic apodeme 7 times longer than paramere; abbreviated streak (Figure 165). ventral sinus inversely acuminate; and marginal DISTRIBUTION (Figure 361).—Known only from denticles less conspicuous (4 specimens examined). northern Panama. Female Genitalia: As in P. latilira. ETYMOLOGY.—The trivial name is a Latin com- VARIATION.—There is no conspicuous variation pound noun formed from the adjective latus (wide) other than that noted for P. latilira. + the noun lira (slide). I refer to the width of the DISTRIBUTION (Figures 360, 361).—The known epipleural fold. range extends from Chiapas, Mexico, to Cartago, LOCALITY RECORDS (Figure 361).—I examined 16 specimens Costa Rica. from Central America. PANAMA: Provincia de Chiriqui: ETYMOLOGY.—The epithet is a Latin compound Bugaba (BMNH, 3 males, holotype, and 1 female; GEKI, 5 males and 1 female; MNHP, 4 males and 1 female). Cor- noun formed from the adjective prolixus (long) + rigimiento de Caldera (GEKI, 1 male). the noun penis (penis). I refer to the length of the aedeagus. REMARKS.—Specimens of this species were mistaken for the superficially similar P. distinctus LOCALITY RECORDS (Figures 360, 361) .—I examined 5 adult (Figure 239), and P. nigriventris. Elytral color specimens from Central America. MEXICO: Estado de Chiapas (GEKI, 1 male). COSTA RICA: Provincia de characteristics of female P. latilira accentuate the Cartago: Turrialba (FMNH, 2 males, holotype, and 1 fe- similarity between that species and the aforemen- male; GEKI, 1 female; ZMAN, 1 male). tioned congeners, but the broad and convex epipleural fold easily segregates P. latilira females REMARKS.—As mentioned in the diagnosis, only from females outside the limbatus group. the differences in male genitalia reliably distinguish P. prolixipenis from P. latilira. I am confident that the noted differences reflect reproductive isolation. 15. Perilypus prolixipenis, new species If the differences were geographic, one would expect FIGURES 172-174, 360, 361 a less than identical appearance of genital characteristics between the P. prolixipenis specimen from TYPE-LOCALITY.—Turrialba, Provincia de Car- Mexico and those from Costa Rica. tago, Costa Rica. Three locality labels have the abbreviation NUMBER 227 59

172

FIGURES 172-180.—Perilypus prolixipennis: 172, tegmen, lateral view; 173, tegmen, ventral view; 174, phallus. P. testaceicornis: 175, habitus; 176, tegmen, lateral view; 177, tegmen, ventral view; 178, antenna; 179, ovipositor, ventral view; 180, ovipositor, dorsal view. (Scale = 1 mm) 60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY "acq," which, I suspect, refers to the Latin word Front Tarsal Claws: Feebly asymmetrical in acquisitus (to add to, to obtain). I suspect that both sexes. Hayne used this abbreviation to identify the col- Male Genitalia (Figures 176, 177): Strongly pig- lector in the same way that one would use "leg" mented. Combined length of phallobase and phallo- (abbreviation for legulns meaning a gatherer, a basic apodeme 6.5 times longer than paramere. collector). One specimen has two different locality Combined length of paramere and phallobase 3.4 records. One of these indicates that the specimen times longer than phallobasic apodeme. Paramere: originates from "Guatemala, Chiapas," the other apex slightly declinate; dorsum narrow, feebly label confines the locality to "Chiapas," which was convex; venter feebly concave; mesoventral margin once part of Guatemala. sinuous; mesodorsal margin concave. Sinus: elliptical dorsal narrower than lanceolate ventral. 16. Perilypus testaceicornis (Pic), Phallus: phallic plates acuminate; marginal den- new combination ticles shallow, broad, and restricted to middle region of phallic plates (3 specimens examined). FIGURES 175-180, 361 Female Genitalia (Figures 179, 180): Dorsal Colyphus testaceicornis Pic, 1941:11 [holotype: female, de- lamina convex laterally, incision short; lateral posited in MNHP; type-locality: Panama]. lobes of ventral lamina shorter than medial lobe; DIAGNOSIS.—Specimens of this species are easily proctigeral bacculi fused as in Figure 180, coxite recognized by the following combination of femoral with small coxital plate (1 specimen examined). color; front and middle femora testaceous, hind VARIATION.—Structural: The specimens examined femur black in distal half, testaceous in proximal do not vary appreciably in external characteristics. half. Color: The sutural, apical, and epipleural DESCRIPTION.—Form (Figure 175): Shallow regions of the elytron may be dark brown or black. bodied; elytra feebly concave postmedially. The paler, nearly fulvous coloration of the elytral Size: Length: males, average about 7.5 mm, range disc suggests a dimly defined vitta that varies in 6.6-8.2 mm; females, average about 7.3 mm, range width. In general, lighter elytra are reddish, darker 6.0-8.7 mm. Width: males, average about 2.0 mm, ones purpurescent. range 1.8-2.1 mm; females, average about 2.1 mm, NATURAL HISTORY.—In May, near Las Lagunas, range 1.8-2.5 mm. Three males and 10 females Panama, at an altitude of 1400 meters, I collected measured. one female specimen by beating tree foliage. G. C. Color: Cranium, antenna, and prothorax flavo- Champion collected 23 adult specimens on Volcan testaceous, except pronotum sometimes dusky mid- de Chiriqui at elevations of from 920 to 1227 apically and pronotal collar black at sides; legs meters. flavotestaceous, except hind femur black in distal DISTRIBUTION (Figure 361).—To my knowledge half; meso- and metasternum mostly black, former these beetles have only been collected on the fulvous medially; elytron and abdominal sterna Pacific slope of the Cordillera de Talamanca, in variable (see "Variation"). northern Panama. Head: As in P. limbatus except antenna (Figure 178) proportionately longer (AL/PL, 1.9); antennal LOCALITY RECORDS (Figure 361).—I examined 25 adult articles 9 and 10 not as wide as long; and length/ specimens from Central America. PANAMA: Provincia de Chiriqui: Volcan de Chiriqui (AMNH, 1 female; BMNH, 1 width ratio of each male antennal article 2.2:1.4: male and 6 females; GEKI, 1 male and 6 females; ISNB, 1 1.9:1.9:1.8:1.9:1.7:1.5:1.4:1.3:1.9. female; MNHP, 3 females; MNHB, 1 female; RNHN, 1 Pronotum: As in P. limbatus except less convex, female; NMNH, 1 male and 1 female); Las Lagunas (GEKI, subapical depression more sinuous and feebly im- 1 female). "Panama" (MNHP, 1 female, holotype). pressed medially, and side margins of pronotum proper less arcuate. 17. Perilypus decoris, new species Elytron: Posthumeral margin, surface, and epipleural fold as in P. limbatus; otherwise shal- FIGURES 1, 2, 181-185, 187, 188, 361 lower, feebly concave postmedially, and generally TYPE-LOCALITY.—Cerro Campana, Provincia de more slender. Panama, Panama. NUMBER Zil 61

TYPE-SPECIMENS.—The holotype male, deposited apex uncinate; lateral depression shallow and in NMNH, was collected by T. L. Erwin in 1974. elongate; dorsum feebly convex, explanate mesally; The allotype is deposited in GEKI. Three para- venter narrow and concave; mesoventral margin types deposited in repositories noted under "Local- feebly arcuate except apically where strongly re- ity Records." curved; mesodorsal margin feebly sinuous and more DIAGNOSIS.—The right-angled incurvature of the sclerotized at base. Sinus: dorsal narrow, slightly elytral discal vitta (Figure 2), near the elytral apex, expanded at basal half and longer than lanceolate easily distinguishes members of this species from ventral sinus. Phallus: phallic plate feebly sinuous; congeners. The general shape of the elytral discal marginal denticles conspicuous, not extended be- vittae in P. decoris is roughly similar in specimens yond anterior limit of phallic plicae (1 male ex- of P. floralis (Figure 342) of the ornaticollis group, amined). but in P. floralis specimens the vitta is not acutely Female Genitalia (Figures 189, 190): Dorsal lam- incurved and the elytra is more oval than rec- ina deeply incised; ventral lamina reduced to one tangulate. very short triangular lobe; proctigeral bacculi fused DESCRIPTION.—Form: As in Figure 2. as in Figure 190; baccular accumination arcuate Size: Length: males, 7.6; females, average about and projecting anteriorly; coxites slender; coxital 8.8, range 8.3-9.3 mm. Width: males, 2.2; females, plates prominent (2 specimens examined). average about 2.3, range 2.5-2.8 mm. Two males Internal Reproductive Organs: Male (Figures and 3 females measured. 187, 188), anterior accessory glands uniramous, Color: Labrum, clypeus, frons, and cranial venter tightly coiled; posterior accessory glands biramous, flavotestaceous, remainder of cranium black; an- outer branch almost twice as long as inner branch; tenna black, except articles 9 to 11 flavotestaceous; testis composed of 12 follicles. Female as in Figure prothorax predominantly flavotestaceous, with mid- 55 (1 male and 2 females examined). apical, midbasal, and basolateral black maculae; VARIATION.—Structural: The specimens studied femur predominantly flavotestaceous, dark brown at showed no noteworthy structural variation. dorsopreapical half; tiba flavotestaceous or dark Color: The posterior junction between the ely- brown; meso- and metasternum black; elytron pre- tral vittae is less distinct in male specimens. dominantly black, with two white vittae that con- NATURAL HISTORY.—Specimens were collected verge near elytral apex; discal vitta acutely from the type-locality (850 meters) in May, June, incurved to sutural margin apically; abdominal and July. Three adults were collected during night venter black in male, predominantly black in by beating "slash and burn" vegetation; T. L. Er- female (first five visible sterna yellow along pos- win, C. W. O'Brien, and B. Marshall. terior margin). DISTRIBUTION.—Known only from the type- Head: As in P. limbatus except interocular im- locality. pressions (Figure 1) broader and more deeply ETYMOLOGY.—Latin, the adjective decoris (orna- impressed, frontal umbo more prominent, antenna mented); with regard to the striking coloration of as in Figure 181. the elytra. Pronotum: As in P. limbatus except more trans- LOCALITY RECORDS (Figure 361).—I examined 5 adult spec- verse (PL/PW, average about 0.86, range 0.88- imens from Central America. PANAMA: Provincia de Pan- 0.90; 5 specimens measured). ama: Cerro Campana (GEKI, 1 male and 2 females; NMNH, Elytron: As in P. limbatus except epipleural fold 1 male, holotype; WBIV, 1 female). broader and more convex, and EL/EW 4.6 (5 speci- REMARK.—The male paratype is without abdo- mens measured). men. Abdomen: Male pygidium as in Figure 185. Front Tarsal Claws: As in P. limbatus. Male Genitalia (Figures 181-184): Strongly pig- The reventazon Group mented. Combined length of phallobase and phallobasic apodeme 5.1 times longer than paramere. Specimens of the reventazon group are character- Combined length of paramere and phallobase 2.3 ized by having the elytral apical slope acute (Fig- times longer than phallobasic apodeme. Paramere: ures 29-31) and the curvature of the epipleural SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 62

FIGURES 181-192.—Perilypus decoris: 181, antenna; 182, tegmen, lateral view; 183, tegmen, ventral view; 184, phallus; 185, male pygidium; 187, male internal reproductive organs, inner lateral view; 188, male internal reproductive organs, outer lateral view; 189, ovipositor, ventral view; 190, ovipositor, dorsal view. P. reventazon: 186, habitus. P. nigriventris: 191, tegmen, lateral view; 192, tegmen, ventral view. (Scale = 1 mm) NUMBER 227 63

margin at elytral apical fourth acutely arcuate (ex- ured); subapical and prebasal depressions deeply cept in P. orthopleurid.es where both characteristics impressed, former strongly sinuous; pronotal arch are secondarily lost). Within the group there is with few coarse punctations; side margins of pro- little nongenitalic structural variation, but color notum proper boldly convex; fovea small, spheroid, polymorphism is abundant at intraspecific level. and deeply impressed; collar prominent. Color differences usually pertain to the elytral sur- Mesoscutellum: Subquadrate. face and are most often correlated with sex. Elytron: Epipleural margin straight in basal The composite distribution of the 17 species that three-fourths, acutely arcuate in apical fourth; currently comprise this group extends from Mexico apical slope acute; surface arenose; disc deeply im- to Peru. pressed with large punctations; epipleural fold plane; EL/EW, average about 4.8, range 4.4-5.3 (20 specimens measured). 18. Perilypus reventazon, new species Front Tarsal Claws: Males, distinctly asymmetri- FIGURES 3-26, 28, 29, 32-46, 48-55, 186, 363 cal. Female, feebly asymmetrical. Male Genitalia (Figures 43-46): Strongly pig- TYPE-LOCALITY.—Rio Reventazon, Turrialba, mented. Combined length of phallobase and phallo- Estado de Cartago, Costa Rica. basic apodeme 5.5 times longer than paramere. Com- TYPE-SPECIMENS.—The holotype male and the bined length of paramere and phallobase 2.4 times allotype, deposited in NMNH, were collected by longer than phallobasic apodeme. Paramere: apex me in 1973. Sixty-eight paratypes deposited in re- feebly uncinate; lateral depression feebly impressed; positories as noted under "Locality Records." dorsum plane; venter slightly concave; mesoventral DIAGNOSIS.—Similar to P. claudus, P. sapientis, margin concave; mesodorsal margin sinuous and and P. cultratus. Specimens of P. reventazon are conspicuously more sclerotized in basal third than substantially larger than members of P. claudus in apical two-thirds. Sinus: dorsal digitiform; ven- and P. sapientis. From P. cultratus, P. reventazon tral lanceolate. Phallus: marginal denticles extended differs by the less-developed antennal serration, from basal half to plicae (30 specimens examined). more convex pronotum, and shallower interocular Female Genitalia: See page 18. depressions. Internal Reproductive Organs: See page 20. DESCRIPTION.—Form: As in Figure 186. VARIATION.—Structural: Except for the sex di- Size: Length: males, average about 8.4 mm, range morphism present in the front tarsal claws, the 7.2-10.0 mm; females, average about 9.0 mm, range specimens studied are structurally homogeneous 8.0-10.5 mm. Width: males, average about 2.3 mm, externally. range 2.0-2-7 mm; females, average about 2.4 mm, Color: There is substantial color variation and range 1.6-3.0 mm. Ten males and 10 females meas- except for the elytral and abdominal coloration, ured. color polymorphism is not sex correlated. The en- Color: Cranium, pronotum, elytron, and abdo- tire spectrum of color variation presented below is men variable (see "Variation"); antenna black; evident in a series of 45 specimens collected from meso- and metasternum dark brown; femur pre- the type-locality. dominantly flavotestaceous, infuscated dorsoapic- The cranium may "be variously bicolored, but ally; tibia and tarsus brown. usually is predominantly black. If predominantly Head: Interocular depressions deeply impressed, black the cranium is flavotestaceous in the postoc- feebly oblique, and not confluent posteriorly; fron- ular epicranial region. When the flavotestaceous tal umbo prominent; eyes boldly convex; HW/ macula of this region is prolonged anteriorly (along IOW, average about 2.2, range 2.1-2.3 (20 speci- the interocular margin of the eyes) it marks the mens measured); antenna (Figure 8) serrate; AL/ epicranium with a subquadrate black macula; the PLS about 1.9; length/width ratio of each male latter combination of cranial color is most often antennal article 2.2:1.1:1.6:1.8:1.7:1.3:1.5:1.3:1.1: found in males with vittate pronotum. In some 1.2:1.7. males, the black epicranial macula is reduced to Pronotum: Moderately transverse (PL/PW, aver- the extent that the cranium is predominantly flavo- age about 0.88, range 0.83-0.93; 20 specimens meas- testaceous to rufescent. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 64 The pronotum is either flavotestaceous or refus- and 1 female; BMNH, 1 male and 1 female; CASC, 1 male and 1 female; CISC, 1 male; CNCC, 1 male; FMNH, 1 male; cent, and vittate or avittate; its lower sides may GEKI, 18 males and 14 females; CMPP, 1 male and 1 fe- be infuscated or entirely black. The pronotal disc male; UCMV, 1 male; MCZC, 1 male and 1 female; MNHB, is either marked by a midapical black macula or 1 male; MNHP, 1 male and 1 female; MZSP, 1 male and 1 by a black vitta. female; NMNH, 2 males, holotype, and 1 female; ZMAN, 1 Elytral coloration distinguishes an avittate phe- male); Turrialba (CNCC, 1 female; FMNH, 8 females; MNHB, 1 female; NMNH, 1 male and 1 female; ZMAN, non, a univittate phenon, and a bivittate phenon. 1 female. Locality not known: FMNH, 1 male and 1 fe- In an avittate beetle the elytron is entirely black, male. except for a small pale macula at the base. This macula corresponds, in position, to the basal limit of a fully developed vitta. In beetles of the univit- 19. Perilypus nigriventris (Gorham), tate phenon the elytron has a discal vitta that be- new combination gins at the base, gradually obliques toward the su- FIGURES 191, 192, 198, 199, 364 ture, and incurves sharply at its apical limit; the incurved portion may or may not reach the sutural Colyphus nigriventris Gorham, 1886:336 [lectotype: male, de- margin. In members of the bivittate phenon the posited in BMNH, here designated; type-locality: El Zapote, Departamento de Escuintla, Guatemala].—Wolcott, 1927a: elytron has two vitta, one on the epipleural margin S3. and one on the disc. The marginal vitta, which is narrower than the discal one, usually terminates DIAGNOSIS.—Within the reventazon group male at the elytral apical eight. Of 32 male specimens specimens of P. nigriventris are superficially very examined only three have avittate elytra; in one of similar to those of P. latilobus, P. calcaris, and P. these the discal vitta diminishes in the basal half. exilis. Among the four species, however, only P. Female specimens whose pronotum is vittate have nigriventris has symmetrical male front tarsal univittate elytra whereas those that have the prono- claws. Females of these species are more difficult to tum maculated have bivattate elytra; in one female separate. specimen the elytron is entirely flavotestaceous. DESCRIPTION.—Form: As in P. reventazon (Figure The male abdomen is black. In females, visible 186) except more robust. abdominal sterna i to v are either entirely flavo- Size: Length: males, average about 7.5 mm, range testaceous or predominantly brown (when mostly 6.6-8.5 mm; females, average about 8.3 mm, range brown their middle and posterior margin is pale), 7.6-9.4 mm. Width: males, average about 2.0 mm, and vi is entirely brown. range 1.8-2.2 mm; females, average about 2.3 mm, NATURAL HISTORY.—I collected 45 specimens of range 2.2-2.6 mm. Four males and five females this species, from May to July, in Turrialba, Costa measured. Rica. Thirty-eight of these were collected by beat- Color: Clypeus, frons, cranium venter, and post- ing tree branches intertwined with lianas, seven by ocular region of epicranium flavotestaceous; re- sweeping herbaceous hedge growth. All specimens mainder of epicranium and gena black; antenna were collected from a forested ravine adjacent to black; pronotum broadly flavotestaceous dorsolat- Rio Reventaz6n; the altitude of the site is 610 me- erally, piceous ventrolaterally and discally; prono- ters. A. Hayne collected two specimens from Tur- tal collar piceous; prosternum flavotestaceous, meso- rialba at 900 meters. and metasternum piceous; femora flavotestaceous, In-captivity feeding behavior of adult specimens except distal region infuscated; tibia and tarsus is similar to that observed in P. limbatus (Gorham). piceous; elytron piceous, with or without testaceous DISTRIBUTION (Figure 363).—Known only from discal vitta; abdomen entirely black, or partially the type-locality. dark brown and partially flavotestaceous. ETYMOLOGY.—The trivial name, reventazon, con- Head: As in P. reventazon except antenna sex stitutes a noun in apposition and refers to the dimorphic in length (AL/PLS, male 2.1; female type-locality. 1.8) and in proportion of length/width ratio of LOCALITY RECORDS (Figure 363).—I examined 71 adult each antennal article, male 2.0:1.0; 1.5:1.7:1.6: specimens from Central America. COSTA RICA: Estado 1.7:1.7:1.7:1.6:1.3:1.9; female 2.0:1.0:1.0:1.0:1.0:1.0: de Cartago: Turrialba, at Rio Reventazon (AMNH, 1 male 1.0:1.0:1.2:1.0:1.7. NUMBER 227 65

Pronotum: As in P. reventazon except subapical 1 female). COSTA RICA: Provincia de Cartago: Turrialba depression shallower, pronotal arch less convex, and (GEKI, 1 male). pronotum proper side margins less arcuate. REMARKS.—Two P. nigriventris females formed Elytron: As in P. reventazon. part of the syntype series of P. bilineatus, a third Front Tarsal Claws: Symmetrical in both sexes. was identified as P. distinctus. Wolcott's (1927a:33) Male Genitalia (Figures 191, 192): Strongly pig- treatise of this species refers to specimens of P. mented. Combined length of phallobase and phallo- reventazon and P. acus. basic apodeme 6 times longer than paramere. Com- bined length of paramere and phallobase 2.8 times longer than phallobasic apodeme. Paramere: ar- 20. Perilypus sapientis, new species cuate and acuminate; lateral depression shallow FICURES 195-196, 363 and elongate; dorsum plane; venter feebly concave; mesoventral and mesodorsal margin concave. TYPE-LOCALITY.—Montes de Oca, San Pedro, Sinus: dorsal and ventral equal in length and Provincia de San Jos£, Costa Rica. lanceolate. Phallus: feebly sinuous; marginal den- TYPE-SPECIMENS.—The holotype male and the ticles distributed from basal fourth to apical fourth allotype are deposited in NMNH. These and 11 of phallic plate (18 specimens examined). paratypes were collected by C. H. Ballou, 10 in Female Genitalia (Figures 198, 199): Dorsal lam- 1936 and one in 1937. Twenty-six paratypes de- ina bilobed, outer margins arcuate, ventral lam- posited in repositories as noted under "Locality ina trilobed, lateral lobes about equal to median Records." lobe; dorsal bacculus not fused; ventral bacculus DIAGNOSIS.—Within the reventazon group speci- acuminate at about posterior third; coxite robust; mens of this species are superficially most similar coxital plates absent (1 specimen examined). to specimens of P. claudus Wolcott. No one ex- VARIATION.—Structural: The male antenna is ternal characteristic will easily separate all members more slender and proportionally shorter than the of these two species, but males are separable by female antenna (see "Head"). Also, in one female genital differences (compare Figures 195, 201), fe- the side margin of the pronotum proper is more males by elytral form; female specimens of P. conspicuously convex and the elytral apical curva- claudus have shorter and more quadrate elytra. ture is less arcuate than in other specimens. DESCRIPTION.—Form: As in Figure 193. Color: The female elytron has a testaceous discal Size: Length: males, average about 6.8 mm, range vitta that is closer to the posthumeral margin than 6.0-7.8 mm; females, average about 7.2 mm, range to the sutural margin. The discal vitta begins pos- 6.7-7.9 mm. Width: males, average about 2.0 mm, teriad to the humeral margin, ends anteriad to the range 1.8-2.3 mm; females, average about 2.2 mm, apex, and varies in width. The male elytron is range 2.0-2.4 mm. Ten males and 10 females meas- avittate. Also in males, the flavotestaceous postocu- ured. lar macula of the epicranium extends anteriorly, Color: Cranium entirely testaceous, or clypeus marking the epicranium with a black subquadrate frons and cranium venter testaceous and epicranium macula; in females, the epicranial postocular mac- and gena black; antenna black; venter and sides of ula is reduced or absent. Color of abdominal ven- pronotum testaceous; pronotal disc testaceous, with ter is also correlated with sex, being entirely black or without midapical infuscation or with broad in males and predominantly flavotestaceous in fe- piceous vitta that narrows abruptly at subapical de- males. pression (Figure 193); pronotal collar black; pro- DISTRIBUTION (Figure 364).—The single locality sternum testaceous; meso- and metasternum black record from Costa Rica extends the known range or piceous; femur flavotestaceous in basal half, piceous in distal half; tibia piceous, distal region of this species from southwestern Guatemala to pale or not; tarsus piceous; elytron piceous, with central Costa Rica. or without discal vitta; elytral discal vitta begins at LOCALITY RECORDS (Figure 364).—I examined 9 adult speci- midbase extends posteriorly equally distant from mens from Central America. GUATEMALA: Departamento suture and posthumeral margin and ends at elytral de Escuintla: El Zapote (BMNH, 1 male, lectotype, and 2 apical eighth, where feebly explanate (Figure 193) females; GEKI, 2 males and 2 females; MNHP, 1 male and SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 66 or not; abdominal venter entirely black, or predom- one other specimen in July on Veronica brachiata inantly dark brown having middle region of visible Bentham. C. Fernandez collected one specimen sternum i to iv flavotestaceous. from San Jose\ in May, between 1000 to 2000 me- Head: As in P. reventazon except antenna pro- ters. portionally shorter (AL/PLS, 1.7) and length/width DISTRIBUTION (Figure 363).—The known range ratio of each antennal article 2.0:1.2:1.3:1.2:1.3: of this species extends from central Guatemala to 1.0:1.3:1.3:1.0:1.0:1.8. central Costa Rica. Pronotum: As in P. reventazon. ETYMOLOGY.—The epithet represents a genitive Elytron: As in P. reventazon except proportion- case of the latin adjective sapiens (wise). I dedicate ally shorter (EL/EW, average about 4.2, range the name of this species to my learned friend 4.1-5.2; 20 specimens measured) and apical curva- George C. Steyskal. ture more acute. LOCALITY RECORDS (Figure 363).—I examined 28 adult Front Tarsal Claws: Symmetrical in both sexes. specimens from Central America. GUATEMALA: Departa- Male Genitalia (Figures 194-196): As in P. nigri- mento de Guatemala: northwestern outskirt of Guatemala ventris except dorsal and ventral sinus broader and City (GEKI, 1 male). COSTA RICA: Provincia de Alajuela: paramere more acutely arcuate in apical region (3 Alajuela (GEKI, 1 female), Provincia de San Jose; San Ignacio (GEKI, 1 male); Farm La Caja, 8 kilometers, west of San specimens examined). Jose (ZMAN, 1 male); San Pedro, Montes de Oca (GEKI, 5 Female Genitalia: As in P. nigriventris. males and 2 females; MHNM, 2 males; NMNH, 2 males, Internal Reproductive Organs: Male and female holotype, and 2 females); San Jose (FMNH, 1 male and 1 as in P. reventazon (1 male and 1 female examined). female; GEKI, 1 male; NMNH, 2 males and 1 female). VARIATION.—Structural: A male specimen from Provincia de Cartago: Turrialba (GEKI, 1 male; NMNH, 1 Turrialba, Costa Rica, is more robust and has a female; ZMAN, 1 male and 2 females). "Cost Rica" (MNHP, 1 female). larger more convex pronotal arch; the side margins of the pronotum proper are broadly arcuate and REMARKS.—The more robust male specimen from the body form is generally more robust. Female Turrialba, heretofore identified as Colyphus nigri- specimens are shorter; their elytral apical curva- ventris, may not be conspecific; but without addi- ture is more acute than it is in males. Depth of tional data I cannot justify its specific distinction the pronotal subapical depression varies indepen- from other specimens of P. sapientes or its con- dent of sex. specificity with P. nigriventris. Color: All males examined have the cranium testaceous. Partial sex-linked color variation is ap- 21. Perilypus claudus (Wolcott) parent in coloration of the pronotal disc: vittate in 8 of 11 females studied, avittate in all males ex- FIGURES 197, 200-203, 364 amined. Color of elytron and abdominal venter is Derestenus claudus Wolcott, 1927a:32 [holotype: male, de- invariably sex linked: always vittate discally in fe- posited in FMNH; type-locality: Turrialba, Estado de males (in one female, from the type-locality, this Cartago, Costa Rica.]. vitta is faintly visible), avittate in males. Male ab- Cleronomus nigrifrons Chevrolat var. flavicornis Wolcott, dominal sterna are black; those of females are part 1927a:36 [holotype: female, deposited in FMNH; new synonymy]. piceous and part testaceous, except for the last visible sternum, which is totally black. Chevrolat's type-specimen is a color variant of NATURAL HISTORY.—Adult specimens were col- P. claudus. lected in Costa Rica from February to July. I col- DIAGNOSIS.—Males of this species can be reliably lected one female specimen in May by beating tree distinguished from males of other species of Peri- foliage in Alajuela. In June, in Montes de Oca, lypus only by genitalic characteristics as described once a forested environ of San Jos6 and now occu- (see "Male Genitalia") and depicted in Figures pied by the campus of the University of Costa Rica, 200-202. No one characteristic will diagnose P. C. H. Ballon collected 10 specimens on Hamelia claudus females but the combination of small size, erecta Jacquin (probably equals H. patens Jacquin), slender body, and distal confluence of the elytral one specimen on Trichilia havanensis Jacquin, and discal vittae will generally separate them from fe- one on Mirabilis jalapa Linnaeus. Ballou collected males of other Costa Rican congeners. NUMBER 227 67

The shape and position of the discal vitta in P. testis composed of 12 follicles. Female (Figure 55), claudus specimens from Volcan Turrialba is very ovary composed of 12 ovarioles (3 males and 2 similar to that in females of P. sapientis. However, females examined). female P. sapientis is more robust and more quad- VARIATION.—Structural: Elytral punctations are rate in body form than female P. claudus. The slightly shallower in specimen from Volcan Irazu configuration of the discal elytral vitta in P. claudus and in those from Quebrada Florida. females from Monteverde is the same as in females Color: On the basis of leg color, specimens from of P. reventazon (Figure 186), but specimens of the Volcan Irazu and those from Quebrada Florida are latter species are considerably larger than those of assigned to a brown-legged morph. These specimens the former. are considerably homogeneous in color unlike in DESCRIPTION.—Form: As in P. sapientis (Figure other population samples of this species. Color 193) except more slender. characteristics of specimens of the dark-legged Size: Length: males, average about 6.2 mm, range phenon are as follows: Head: cranium testaceous, 5.3-7.3 mm; females, average about 6.8 mm, range immaculate; antenna piceous with articles 9 to 11 5.7-8.0 mm. Width: males, average about 1.7 mm, not testaceous beneath. Thorax: prothorax pre- range 1.4-1.9 mm; females, average about 1.8, dominantly testaceous; pronotum with midapical range 1.7-2.3 mm. Ten males and 10 females macula, or with or without discal vitta, latter inter- measured. rupted at middle or not; pronotal collar piceous; Color: Color characteristics are exceptionally meso- and metasternum piceous; legs piceous; ely- variable and because of this a composite descrip- tron piceous, with cupreous or purpurescent luster. tion of this species' integumental color is not pro- Abdomen: sternum piceous in both sexes. vided. Instead, the general color patterns and the Leg color can also be used to distinguish a yellow- deviations from them are described under "Varia- legged morph. In specimens of this morph the head, tion." antenna, pronotum, mesoscutellum and legs are Head: As in P. reventazon except antenna pro- entirely flavotestaceous, and the elytra purpurescent portionally longer (AL/PLS, 2.0) and length/width and avittate. ratio of each male antennal article 2.8:1.4:1.8:1.8: All other specimens examined exhibit the fol- 1.6:1.6:1.4:1.4:1.2:1.0:2.3. lowing possible color characteristics: Head: cra- Pronotum: As in P. reventazon. nium flavotestaceous,wit h or without midepicranial Elytron: As in P. reventazon. macula, or clypeus and frons flavotestaceous, epi- Front Tarsal Claws: Symmetrical in both sexes. cranium black (except postocular region flavotesta- Male Genitalia (Figures 200-202): Strongly pig- ceous), gena piceous, and venter piceous or mented. Combined length of phallobase and phallo- flavotestaceous; antenna flavotestaceous or black, basic apodeme 5 times longer than paramere. when piceous articles 9 to 11 testaceous or dark Combined length of paramere and phallobase 3 brown. Thorax: pronotum predominantly black or times longer than phallobasic apodeme. Paramere: predominantly flavotestaceous; when predominantly feebly arcuate in lateral view; parameral septum black vittate discally and flavotestaceous laterally; prominent (note arrow in Figure 201); lateral de- when predominantly flavotestaceous discal vitta pression indistinct; dorsum tumid midbasally, feebly reduced; pronotal discal vitta (when present) may convex medially; venter narrow and feebly convex; be entire or interrupted at middle; when pronotum mesoventral margin concave; mesodorsal margin predominantly flavotestaceous midapical macula sinuous. Sinus: dorsal irregular; ventral broadly present; pronotal collar piceous; prosternum elliptical. Phallus: as in Figure 202 (29 specimens flavotestaceous; meso- and metasternum piceous; examined). profemur flavotestaceous in basal half, piceous in Female Genitalia: As in P. nigriventris (2 speci- apical half, meso- and metafemur flavotestaceous in mens examined). basal two-thirds and piceous in apical third, or all Internal Reproductive Organs: Male (Figures femora flavotestaceous in basal half and piceous in 197-203), anterior accessory gland uniramous. apical half; tibia entirely piceous, or piceous in tightly coiled; posterior accessory gland biramous, basal half and testaceous in apical half (usually only outer branch one-fourth longer than inner branch; hind tibia testaceous in apical half); tarsus piceous; SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 68

FIGURES 193-203.—Perilypus sapientis: 193, habitus; 194, tegmen, lateral view; 195, tegmen, ventral view; 196, phallus. P. claudus: 197, male internal reproductive organs; 200, tegmen, lateral view; 201, tegmen, ventral view (arrow indicates parameral septum); 202, phallus; 203, posterior accessory gland. P. nigriventris: 198, ovipositor, ventral view; 199, ovipositor, dorsal view. (Scale = 1 mm) NUMBER 227 69 elytron piceous, with or without purpurescent luster from Mexico extends the known range of this and vittate or avittate discally. Abdomen: venter species from Mexico to Costa Rica. entirely black, or visible sternum i to v flavotesta- LOCALITY RECORDS (Figure 364).—I examined 52 adult ceous infuscated discally and laterally, sternum vi specimens of this species from Central America. MEXICO: piceous. Estado de Morelos: Yautepec (GEKI, 1 male). COSTA RICA: Specimens from Monte Verde are sex dimorphic Provincia de Puntarenas: Monteverde (AMNH, 1 male and 1 with respect to cranial, pronotal, and possibly female; BMNH, 1 male and 1 female; CASC, 1 male and 1 female; FMNH, 1 male and 1 female; GEKI, S males and 6 elytral color. Male specimens have the cranium females; CMPP, 1 male; MNHB, 1 female; MNHP, 1 male; entirely flavotestaceous, pronotal vitta absent, and NMNH, 2 males and 2 females; ZMAN, 1 male and 1 female). elytron avittate. Female specimens have the cranium Provincia de Cartago; Turrialba (FMNH, 2 males, holotype, predominantly piceous, the pronotal vitta is always and 2 females; GEKI, 1 male); Volcan Turrialba (GEKI, 10 entire, and of 14 specimens eight have vittate males and 1 female); Volcan Irazii (FMNH, 1 female; GEKI, 1 female; NMNH, 3 females). Provincia de Limdn: Que- elytra. The elytral discal vitta, which is nearer to brada Florida (GEKI, 2 males; NMNH, 1 male and 1 female). the posthumeral margin than to the sutural margin, is incurved apically; the incurvature reaches the REMARKS.—The vast color variability present in sutural margin. the available samples is evident at intra- and inter- I studied too few specimens from Volcan Turri- populational levels. Diversity of integumental color alfa to determine relationships between color and is particularly evident in coloration of the elytron. sex. The following color characteristics are present, The elytral discal vitta may be linear or incurved however. The legs are entirely flavotestaceous in 5 at the apex. Elytra are avittate in the beetles from of 11 males; in the other males the legs are partly Volcan Irazu and Quebrada Florida. piceous and partly flavotestaceous. Among all speci- While intrapopulation color polymorphism is mens studied, 2 males and the only female from this not extraordinary in Perilypus the extent reported locality have univittate elytra while in another here is more than usual. Accordingly, one might male the elytral vitta is reduced to a short basal suspect that the beetles from Monteverde, Volcan streak. Turrialba, Volcan Irazu, and Quebrada Florida are NATURAL HISTORY.—In June, on the northeastern not conspecific, as indeed they may not be. How- slope of Volcan Turrialba, Costa Rica, at an alti- ever, for the present I assume them conspecific in tude of 1450 meters, I collected a series of 12 speci- view of their structural similarity including char- mens by beating live branches of Psidium guajava acteristics of the genital organs of both sexes. Linnaeus. Wolcott's (1927a:36) description of Derestenus I collected 26 additional specimens, in June, from nigrifrons Chevrolat var. flavicornis is based on one Monte Verde, Costa Rica, by beating dense under- female specimen from Turrialba. Integumental story composed mostly of live and dead tree color (Wolcott's criteria for recognizing var. flavi- branches and liana vines. The macrohabitat of the cornis) of a male specimen from Monte Verde, locality may be described as a dry section of mon- Costa Rica, is identical to that of Wolcott's holo- tane forest situated at 1600 meters on the Pacific type. There is no noticeable anatomical difference between these two specimens except for size and slope of the Cordillera de Tilaran. Holdridge, et al. structures related to sex. I believe the specimens (1971), classified this region as "lower montane rain under consideration are conspecific, not only to each forest." other, but also with the other specimens included In April and May, F. Nevermann collected four herein. I assign two male specimens and Wolcott's specimens of the brown-legged phenon from the var. flavicornis to a yellow-legged phenon (see western slope of Volcan Irazii (at 1500 meters); "Variation)." four others were collected during May near Que- brada Florida (at 1400 meters). A single specimen, with an F. Nevermann label but apparently col- 22. Perilypus cultratus, new species lected by "Schild" (the writing is not clear), was collected in October from Turrialba. FIGURES 204-206, 211, 216, 363 DISTRIBUTION (Figure 364).—The single specimen TYPE-LOCALITY.—Rio Reventazdn, Turrialba, SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 70 Estado de Cartago, Costa Rica. pigmented. Combined length of phallobase and TYPE-SPECIMENS.—The holotype male is depos- phallobasic apodeme 7 times longer than paramere. ited in NMNH, the allotype in GEKI; both were Combined length or paramere and phallobase 3 collected by me in 1971. Five paratypes are depos- times longer than phallobasic apodeme. Paramere: ited in the repositories noted under "Locality acuminate in lateral view; lateral depression not Records." conspicuous; dorsum feebly convex; venter concave; DIAGNOSIS.—Within the reventazon group, speci- mesoventral and mesodorsal margin sinuous, mar- mens from Costa Rica belong to this species if gins confluent in apical half, not in distal half. their pronotal disc is faintly tumid behind the Sinus: dorsal narrow, as long as inversely acuminate subapical depression and in front of the prebasal ventral. Phallus: attenuate; marginal denticles depression, if the pronotal foveae are deeply im- prominent (4 specimens examined). pressed and not particularly spheroid (sometimes Female Genitalia: As in P. nigriventris. they approach a crescentic shape), and if their Internal Reproductive Organs: Male and female antenna is more serrate than in P. reventazon as in P. reventazon (1 male and 2 females exam- (Figure 8). Specimens of this species also can be ined). easily separated from the very similar sympatric VARIATION.—Structural: The interocular depres- specimens of P. reventazon by differences in the sions are less depressed and abbreviated in the male pygidium (compare Figures 211, 39) and by female specimen from Hamburg Farm; they are the lineate female elytral vitta. conspicuously rugose in the male from Tuis and in DESCRIPTION.—Form: As in P. reventazon (Figure one female from Turrialba. 186) except pronotum less convex. Color: The pronotal vitta is deltoid in the female Size: Length: males, average about 7.6 mm, range specimen from Hamburg Farm, and in one from 8.0-6.6 mm; females, average 8.0 mm. Width: males, Turrialba. Except for the male from Hamburg average about 2.2 mm; range 2.0-2.6 mm; females: Farm, male elytra are avittate; female elytra are 2.4 mm. Five males and 2 females examined. vittate discally. Color: Clypeus, frons, postocular region of epi- NATURAL HISTORY.—In June and July, within a cranium, and cranial venter testaceous; gena and forested ravine adjacent to Rio Reventaz6n, I col- remainder of epicranium black; antenna black; side lected a male and two females by sweeping her- margin of pronotum proper flavotestaceous; pro- baceous hedge growth. notal disc broadly vittate; pro-, meso-, and meta- In June, within the Rio Reventaz6n drainage sternum dark brown; femora predominantly system, in Limon, F. Nevermann collected one flavotestaceous, infuscated dorsoapically; elytron female in "troknem holz" (probably translated as piceous, with or without short flavotestaceous discal fallen timber, or from a wood pile). He also col- vitta; abdomen entirely black or predominantly lected a male in August from Hamburg Farm. flavotestaceous. DISTRIBUTION (Figure 363).—The few locality Head: As in P. reventazon except: interocular records suggest that this species is primarily distrib- depressions more deeply impressed, rugose or not; uted along the Costa Rican Atlantic drainage antenna more serrate; and length/width ratio of system. each male antennal article 2.4:1.2:1.4:1.3:1.3:1.3: ETYMOLOGY.—Latin, the adjective cultratus 1.2:1.0:1.2:1.1:2.2. (knife-shaped); the trivial name refers to the knife- Pronotum: As in P. reventazon except foveae shaped phallus. more deeply impressed, more elongate than sphe- LOCALITY RECORDS (Figure 363).—I examined 7 adult spec- roid; and disc behind subapical depression and imens from Central America: COSTA RICA: Provincia de before prebasal depression feebly tumid. Cartago: Turrialba at Rio Reventaz6n (GEKI, 2 females; Elytron: As in P. reventazon. NMNH, 1 male, holotype); Tuis (GMNH, 1 male). Provinda Front Tarsal Claws: Male and female feebly de Lim6n: Hamburg Farm at Rio Reventaz6n (GEKI 1 male; asymmetrical. NMNH, 1 female). Locality not known (FMNH, 1 male). Abdomen: Male pygideal apodeme as long as REMARKS.—Females of this species have been pygidial plate (Figure 211). confused with females of P. bilineatus; however, Male Genitalia (Figures 204-206, 216): Strongly the conspicuous convexity of the elytron and the NUMBER 227 71 distinct confluence of the epipleural and discal nal article 2.5:1.0:1.1:1.4:1.2:1.1:1.1:1.1:1.0:1.0:1.8. vittae at the elytral apex in P. bilineatus should Pronotum: As in P. reventazon except as long as easily separate specimens of that species from those wide, subapical depression more sinuous, and pro- of P. cultratus. notal arch transversely wrinkled midapically. Elytron: As in P. reventazon except shallower and more convex. 23. Perilypus bilineatus (Gorham), Front Tarsal Claws: Male, damaged. Female, new combination symmetrical.

FIGURES 207-209, 364 Male Genitalia (Figures 207-209): Strongly pigmented. Combined length of phallobase and Colyphus bilineatus Gorham, 1886:335 [lectotype: female, de- phallobasic apodeme 6 times longer than paramere. posited in BMNH, two paralectotypes, females, one de- Combined length of paramere and phallobase 2.5 posited in BMNH and one in ZMAN, here designated; type-locality: Volcan de Chiriqui, Provincia de Chiriquf, times longer than phallobasic apodeme. Paramere: Panama].—Wolcott, 1927a:37. feebly uncinate in ventral view; lateral depression deeply impressed; dorsum convex; venter concave; DIAGNOSIS.—The slender, shallow body combined mesoventral margin concave; mesodorsal margin with the following characteristics will distinguish sinuous. Sinus: dorsal ovate; ventral lanceolate. specimens of this species within the reventazon Phallus: phallic plates tapered; marginal denticles group; elytral apex testaceous, elytral disc vittate stout (1 specimen examined). or not, pronotum as long as wide, and pronotal Female Genitalia: As in P. nigriventris. subapical depression shallow and wrinkled trans- VARIATION.—Structural: Antenna is proportion- versely midbasally. ally shorter and more cylindrical in one female DESCRIPTION.—Form: As in P. reventazon (Figure specimen, particularly articles 8 to 10. 186) except shallower and more slender. Color: In the male specimen the pronotum is Size: Length: male, average 6.7 mm; females, predominantly testaceous, the abdomen black, and average about 7.3 mm, range 6.4-8.5 mm. Width: the elytral disc avittate. Females have the pronotum males 2.0; females, average about 2.1 mm, range predominantly black, abdomen part black and part 2.0-2.4 mm. One male and 5 females measured. testaceous, and the elytral disc distinctly vittate. The Color: Cranium entirely testaceous, or clypeus elytral epipleural vitta varies in extent of expres- frons and postocular region of epicranium tes- sion, and is absent in the female specimen in which taceous and remainder of cranium black; antenna antennal articles 8 to 10 are more cylindrical. black; pronotum predominantly testaceous and infuscated midapically, or predominantly black and NATURAL HISTORY.—G. C. Champion collected pronotum proper with side margins and anterior the available specimens from Volcan de Chiriquf at angles testaceous; prosternum testaceous; meso- and various altitudes ranging from 610 to 1829 meters. metasternum black; femur predominantly testa- DISTRIBUTION.—Presently known only from the ceous, infuscated apically; tibia and tarsus dark type-locality. brown; elytron predominantly piceous, epipleuron and apex faintly testaceous and flavotestaceous dis- LOCALITY RECORDS (Figure 364).—I examined 6 adult specimens from Central America. PANAMA: Provincia de Chri- cal vitta closer to posthumeral margin than to qui: Volcan de Chiriqui (BMNH, 1 male and 2 females, lecto- sutural margin (as in Figure 264); discal vitta may type; GEKI, 1 female; MNHP, 1 female; ZMAN, 1 female). or may not be confluent with testaceous epipleural vitta; abdominal venter black, or visible sterna i to REMARKS.—The name of this species has been iv testaceous and infuscated anteriorly, v predomi- applied to members of P. nigriventris, P. reventazon, nantly testaceous or predominantly dark brown, and P. copiscolis. In view of the substantial simi- and vi black. larity in elytral color among females of these species, Head: As in P. reventazon except cranium some- particularly with regard to the form of elytral times conspicuously macrosculptured; antenna less vittae, it is not surprising that the aforementioned serrate; and length/width ratio of each male anten- misidentincations involved only female specimens. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 72

FIGURES 204-216.—Perilypus cultratus: 204, tegmen, lateral view; 205, tegmen, ventral view; 206, phallus; 211, male pygidium; 216, tegmen (arrow indicates confluency between mesoventral and mesodorsal margins). P. bilineatus: 207, tegmen, lateral view; 208, tegmen, ventral view; 209, phallus. P. cerroazul: 210, posterior accessory gland; 212, tegmen, lateral view; 213, tegmen, ventral view; 214, tegmen, dorsal view (arrow indicates dorsal brace); 215, anterior accessory gland. (Scale = 1 mm) NUMBER 227 73

24. Perilypus cerroazul, new species Panama, W. Bivin collected the holotype specimen by beating vegetation at 700 meters. FIGURES 210, 212-215, 217, S64 ETYMOLOGY.—The epithet is a noun in apposition TYPE-LOCALITY.—Cerro Azul, Provincia de Pan- and refers to the type-locality. amd, Panama. LOCALITY RECORDS (Figure 364).—I examined 1 adult speci- TYPE-SPECIMENS.—The holotype male, deposited men from Central America. PANAMA: Provincia de Pan- in NMNH, was collected by W. Bivin in 1971. ama: Cerro Azul (NMNH, 1 male, holotype). DIAGNOSIS.—The only specimen studied is superficially very similar to the sympatric specimens of 25. Perilypus ordinatus, new species P. ordinatus. However, the antenna is more distinctly serrate and the pygidium more robust FIGURES 218-221, 245, 364 (compare Figures 217, 245) in male specimens TYPE-LOCALITY.—Cerro Campana, Provincia de of P. cerroazul. Panama, Panama. DESCRIPTION.—Form: As in P. reventazon (Figure TYPE-SPECIMENS.—The holotype male, deposited 186) except proportionally longer. in NMNH, was collected by W. Bivin in 1971. Five Size: Length: 9.2 mm. Width: 2.4 mm. paratypes deposited in the repositories noted under Color: As in P. reventazon except postocular "Locality Records." region of epicranium broadly flavotestaceous, femur DIAGNOSIS.—See P. cerroazul. black at apex only, and elytron with faint discal DESCRIPTION.—Form: As in P. reventazon (Figure vitta visible only at elytral basal half. 186). Head: As in P. reventazon except antenna more Size: Length: Average about 7.5 mm, range serrate. 6.8-8.5 mm. Width: Average about 2.1 mm, range Pronotum: As in P. reventazon except more 1.8-2.4 mm. Three males measured. convex and side margins of pronotum proper less Color: Clypeus, frons, cranium venter, and arcuate. postocular region of epicranium flavotestaceous; Elytron: As in P. reventazon except proportion- remainder of epicranium and gena black (see ally longer (EL/EW, 5.2) and more densely "Variation"); antenna black; pronotum with side pubescent. margin broadly flavotestaceous, lower sides and Abdomen: Male pygidium (Figure 217) partic- disc black; pronotal collar black (see "Variation"); ularly robust. prosternum testaceous; meso- and metasternum Front Tarsal Claws: Feebly asymmetrical. black; profemur black dorsally, flavotestaceous Male Genitalia (Figures 212-214): Strongly pig- ventrally; meso- and metafemur black in dorso- mented. Combined length of phallobase and apical fourth, flavotestaceous in remainder; pro- phallobasic apodeme 9 times longer than paramere. and mesotibia black, metatibia at least partially Combined length of paramere and phallobase 4 testaceous; elytron piceous; abdomen black. times longer than phallobasic apodeme. Paramere: Head: As in P. reventazon except antenna more acuminate and arcuate in lateral view; lateral serrate, proportionally shorter (AL/PL, 1.7) and depression indistinct; dorsum narrow and feebly frontal umbo more prominent. convex; venter broadly concave; mesodorsal margin Pronotum: As in P. reventazon except less trans- feebly concave, strongly braced, and margins verse (PL/PW, average about 0.94, range 0.92-0.96; contiguous at basal fifth (note arrow in Figure 3 specimens measured), more convex, and side 214). Sinus: dorsal and ventral broadly lanceolate. margins of pronotum proper less arcuate. Phallus: acuminate. Elytron: As in P. reventazon except more densely Internal Reproductive Organs: Male (Figures 210, pubescent. 215), anterior accessory gland uniramous, coiled; Abdomen: Male pygidium as in Figure 245. posterior accessory gland biramous, outer branch Front Tarsal Claws: Feebly asymmetrical. about twice as long as inner branch; testis composed Male Genitalia (Figures 218-220): Strongly pig- of 12 follicles. mented. Combined length of phallobase and NATURAL HISTORY.—In June, on Cerro Azul, phallobasic apodeme 3.5 times longer than para- 74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY mere. Combined length of paramere and phallo- 185 (compare with Figure 245). Female specimens base 3.6 times longer than phallobasic apodeme. of these three species are difficult to separate, but Paramere: lateral depression broad, extended be- color of the abdominal venter seems to provide yond basal limit of paramere; dorsum narrow and some basis for distinguishing them. The abdominal convex; venter narrow and concave; mesodorsal venter is dark brown in the anterior half and margin concave, strongly incised at basal third, flavotestaceous in the posterior half (except the mesoventral margin strongly sinuous, feebly incised fifth visible sternum is infuscated) in females of at basal third and uncinate preapically (note arrow P. calcaris, entirely dark brown in females of P. in Figure 219). Sinus: as in Figure 219. Phallus: as lobulatus, and predominantly flavotestaceous (ster- in Figure 220 (5 specimens examined). num vi is dark brown) in females of P. nigriventris. Internal Reproductive Organs: Male (Figure 221); DESCRIPTION.—Form: As in P. reventazon (Figure anterior accessory gland uniramous, tightly coiled; 186). posterior accessory gland biramous, outer branch Size: Length: males, average about 7.6 mm, range longer than inner branch; testis composed of 12 7.2-7.8 mm; female 7.9 mm. Width: males, average follicles (5 specimens examined). about 2.3 mm, range 2.0-2.4 mm; female, 2.2 mm. VARIATION.—Structural: I found no noteworthy Three males and one female measured. structural variation among the specimens studied. Color: As in P. nigriventris except female elytron Color: In one specimen the head, pronotum, and faintly pale along posthumeral margin, femora dark mesoscutellum is entirely flavotestaceous. brown in apical half (except testaceous ventroapi- NATURAL HISTORY.—On Cerro Campana (850 cally), coxae strongly infuscated, and female abdom- meters) W. Bivin collected two specimens by beat- inal venter not predominantly flavotestaceous. ing, one during March and one during May. From Head: As in P. reventazon except antenna more the same locality additional specimens were slender. collected by S. S. and W. D. Duckworth (in April Pronotum: As in P. reventazon except pronotal by sweeping) and by H. P. Stockwell (in May by arch and pronotum proper with shallow rugosities. beating). Elytron: As in P. reventazon except punctations ETYMOLOGY.—Latin, the adjective ordinatus (well smaller and shallower and surface more arenose. ordered). I dedicate the name of this species to my Front Tarsal Claws: Male, distinctly asymmet- colleague and friend Terry L. Erwin. rical; female, feebly asymmetrical. LOCALITY RECORDS (Figure 364).—I examined 6 adult speci- Male Genitalia (Figures 222-224): Strongly pig- mens from Central America. PANAMA: Provincia de Pan- mented. Combined length of phallobase and phal- ama: Cerro Campana (GEKI, 2 males; HPST, 1 male; lobasic apodeme 5.3 times longer than paramere. NMNH, 2 males, holotype; WBIV, 1 male). Combined length of paramere and phallobase 4 times longer than phallobasic apodeme. Paramere: 26. Perilypus calcaris, new species apex feebly uncinate in ventral view; lateral depression narrow, more dorsad than ventrad; FIGURES 222-226, 363 dorsum convex and broadly explanate medially; TYPE-LOCALITY.—Finca Recreo, Municipio de venter narrow and feebly concave; mesoventral Yepocapa, Departamento de Chimaltenango, Gua- margin emarginated and biacuminate; mesodorsal temala. margin sinuous, with stout brace at base. Sinus: TYPE-SPECIMENS.—The holotype male and the dorsal much narrower than elliptical ventral. allotype, deposited in FMNH, were collected by Phallus: acuminate; outer margin inflexed; stout R. D. Mitchell in 1948. Two paratypes: BMNH, marginal denticles centralized at middle half of 1 male; GEKI, 1 male. phallic plate (3 specimens examined). DIAGNOSIS.—Specimens of P. calcaris are super- Female Genitalia (Figures 225, 226): As in P. fically very similar to specimens of P. nigriventris orthopleuridus except dorsal lamina reduced. and of P. lobulatus. The pygidium, however, is VARIATION.—Structural: The pronotal rugosities substantially larger in males of P. calcaris, being vary in extent of expression. about equal in size to the one illustrated in Figure Color: Color of the abdominal venter varies with NUMBER 227 75

FIGURES 217-228.—Perilypus cerroazul: 217, male pygidium. P. ordinatus: 218, tegmen, lateral view; 219, tegmen, ventral view (arrow indicates preapical uncination of mesoventral margin); 220, phallus; 221, male internal reproductive organs. P. calcaris: 222, tegmen, lateral view; 223, tegmen, ventral view (arrow indicates acumination of mesoventral margin); 224, phallus; 225, ovipositor, ventral view; 226, ovipositor, dorsal view. P. orthopleuridus: 221, ovipositor, ventral view; 228, ovipositor, dorsal view. (Scale = 1 mm) 76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY sex; in male specimens entirely dark brown, in range 2.0-2.8 mm; females average about 2.3 mm, females dark brown in basal half and mostly range 2.0-3.4 mm. Ten males and 10 females flavotestaceous in apical half. measured. NATURAL HISTORY.—R. D. Mitchell collected Color: Cranium unicolorous or bicolorous; uni- three adult specimens (at 1341 meters) in May from colorous cranium is flavotestaceous; bicolorous Finca Recreo, Guatemala, by beating "dead cranium has clypeus, frons, and cranial venter branches." G. C. Champion collected one specimen flavotestaceous, and epicranium and gena piceous; from Cerro Zunil, Guatemala, at an elevation antenna entirely flavotestaceous, or articles 1 to 8 between 1219 and 1832 meters. brunneus and 9 to 11 flavotestaceous; pronotum DISTRIBUTION (Figure 363).—Known only from predominantly flavotestaceous and with midapical the southwestern highlands of Guatemala. piceous macula, or flavotestaceous at sides and piceous discally; pronotal collar piceous or pre- ETYMOLOGY.—The trivial name is the genetive dominatly flavotestaceous; prosternum flavotestace- case of calcar (a spur) and refers to the spur-like ous; mesoscutellum and meso- and metasternum acumination on the mesoventral margin of para- flavotestaceous or piceous; femur entirely flavo- meres. testaceous, flavotestaceous in basal half and ventro- LOCALITY RECORDS (Figure 363).—I examined 4 adult speci- apical half and piceous in dorsoapical half, or mens from Central America. GUATEMALA: Departamento predominantly flavotestaceous and infuscated dorso- de Quetzaltenango: Cerro Zunil (BMNH, 1 male). Depart- apically; tibia and tarsus testaceous or piceous; amento de Chimaltenango: Finca Recreo (GEKI, 1 male; elytron piceous, with or without flavous discal FMNH, 1 male holotype, and 1 female). vitta; visible abdominal sterna I to v entirely flavotestaceous, predominantly flavotestaceous and 27. Perilypus orthopleuridus (Thomson), infuscated basally or laterally, or piceous; sixth new combination visible abdominal sternum piceous. Head: As in P. reventazon except antenna less FIGURES 27, 227-234, 364 serrate. Derestenus orthopleuridus Thomson, 1860:57 [holotype, fe- Pronotum: As in P. reventazon except more male, deposited in MNHP; type-locality: Mexico].—Gor- convex. ham, 1882:144—Schenkling, 1906:263. Elytron: As in P. reventazon, except apical slope Derestenus vittipennis Chevrolat, 1874:290 [lectotype: male, deposited in BMNH, here designated; type-locality: Mex- gradual, punctations shallower, surface not arenose, ico; new synonmy].—Gorham, 1882:143. and female epipleuron explanate and plane ven- Derestenus collaris Chevrolat, 1876:12 [lectotype: female, de- trally (Figures 230, 231). posited in MNHP, here designated; type-locality: Mexico]. Abdomen: Male pygidium (Figure 229) con- —Schenkling, 1906:263. spicuously large and feebly convex. The new synonmy is based on the discovery that Front Tarsal Claws: Male, boldly asymmetrical; D. vittipennis Chevrolat is the heretofore unde- female, feebly asymmetrical. scribed male of P. orthopleuridus. Male Genitalia (Figures 232-234): Strongly pig- DIAGNOSIS.—The combination of front tarsal mented. Combined length of phallobase and claws boldly asymmetrical, and antenna partially or phallobasic apodeme 4.7 times longer than para- totally flavotestaceous will easily separate males of mere. Combined length of paramere and phallobase this species from other males within the reventazon 3.0 times longer than phallobasic apodeme. Para- group. The epipleuron (Figure 230) is explanate mere: apex uncinate; lateral depressions extended and plane ventrally in P. orthopleuridus females beyond base of ventral sinus; dorsum particularly only. narrow and broadly braced basally; venter narrow, DESCRIPTION.—Form: As in P. reventazon (Figure concave; mesoventral margin emarginated (note 186) except elytral apical slope gradual. arrow in Figure 233); mesodorsal margin sinuous. Size: Length: males, average about 8.2 mm, range Sinus: dorsal about twice as broad as ventral. 5.8-9.9 mm; females, average about 8.9 mm, range Phallus: acuminate; outer margin inflexed in basal 7.6-10.4 mm. Width: males, average about 2.1 mm, half and denticulated from middle to basal three- NUMBER 227 77 fourths; plicae feebly developed (45 specimens Female Genitalia (Figures 227, 228): Dorsal and examined). ventral lamina and coxites very long; dorsal lamina

FIGURE 229-238.—Perilypus orthopleuridus: 229, male pygidium, 230, female elytron, ventral view, X 25 (arrow indicates epipleuron); 231, female elytron, lateral view, X 25 (arrow indicates epipleuron); 232, tegmen, lateral view; 233, tegmen, ventral view (arrow indicates acumination of mesoventral margin); 234, phallus, lateral view. P. exilis: 235, tegmen, lateral view; 236, tegmen, ventral view. P. latilobus: 237, tegmen, lateral view; 238, tegmen, ventral view. (Scale = 1 mm) 78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY slender, deeply incised; lateral lobes of ventral males and 3 females; GEKI, 2 males; MNHB, 1 female, lamina shorter than medial lobe; inner margin of LACM, 1 male; MNHP, 3 females, holotype; ZMAN, 1 male); 1 mile [1.6 km] west of Fortin de las Flores (CISC, 1 male); coxites well sclerotized; remainder as in P. reventa- Cordoba (BMNH, 2 males; CASC, 4 males and 2 females; zon (2 specimens examined). FMNH, 1 male; GEKI, 4 males and 2 females; NMNH, 2 Internal Reproductive Organs: Male, anterior males); Jalapa (CASC, 1 male; FMNH 1 male); Mirador accessory gland uniramous tightly coiled, posterior (MNHB, 1 male); Lago de Catemaco (CNCC, 1 male; GEKI, accessory gland biramous (3 specimens examined). 2 males and 1 female); San Andres Tuxtla (GEKI, 1 male); Playa Vincente (BMNH, 1 female); Cerro de Palmas (BMNH, VARIATION.—Structural: The epipleuron is ex- 1 female). Estado de Oaxaca: Oaxaca (BMNH, 1 female, planate in female elytra only; epipleural explana- GEKI, 1 female); Temazcal (GEKI, 1 female; GHNE, 1 fe- tion varies but is always prominent. The cranium, male). Estado de Chiapas: 8 miles [13 km] south of Simojovel femora, pronotum, and abdominal sterna are (CNCC, 1 male); 27 kilometers southwest of Simojovel (GEKI, conspicuously rugose and the elytral surface is more 1 female); 2 miles [3 km] northeast of Bochil (GEKI, 1 male); 2 kilometers north of Bochil (GEKI, 1 female); 12 miles [19 arenose in the female specimen from Temazcal, km] north of Tuxtla Gutierrez (GEKI, 1 male); 11 miles [18 Mexico. km] north of Ocozocoautla (GEKI, 1 female; TAMT, 1 fe- Color: Coloration of cranium, pronotum, femur, male); 22 kilometers north of Ocozocoautla (CNCC, 1 female); meso- and metasternum, and of abdominal sterna El Suspiro, Berriozabal (GEKI, 1 male). "Mexico" (BMNH, 1 is correlated with sex. male and 2 females; FMNH, 1 male; GEKI, 3 males and 7 females; MNHB, 1 female; MNHP, 1 male and 7 females; Pronotal color distinguishes 2 male phena, a ZMAN, 2 females). GUATEMALA: Departamento de Alta maculate phenon and a vittate phenon. Among the Verapaz: San Juan (BMNH, 1 male). Tamahu (AMNH, 1 45 male specimens studied, 35 belong to the female); Senahii (BMNH, 1 female). HONDURAS: Depart- maculate phenon (the pronotum is predominantly amento de Choluteca: Cantarranas, Rio Choluteca (GEKI, 1 flavotestaceous and has a piceous midapical macula) female.) COSTA RICA: Provincia de Cartago: Turrialba and 10 to the vittate phenon (the pronotal disc (GEKI, 1 male). has a piceous vitta that is narrowly percurrent or REMARKS.—Specimens of this species were iden- interrupted at the middle). tified as P. quadrilineatus, P. distinctus, P. nigri- Among the maculate males, 11 have the elytron frons, and Colyphus signoticollis Spinola. distinctly vittate, in the remaining specimens the elytral vitta is faintly visible or absent. In all 28. Perilypus exilis, new species maculate males the cranium, femur, and meso- and metasternum are entirely flavotestaceous. In two of FIGURES 235, 236, 363 the 10 vittate males the elytra are distinctly vittate, TYPE-LOCALITY.—Thirty-two kilometers north of while in the remaining 8 the elytral discal vitta Huixtla, Estado de Chiapas, Mexico. are either faintly visible or absent. Vittate males TYPE-SPECIMENS.—The holotype male, deposited have the cranium and the femur bicolorous, and in CNCC, was collected by W. R. M. Mason in the meso- and metasternum piceous. In all males 1969. the abdomen is entirely piceous. DIAGNOSIS.—The combination of conspicuously The female pronotum is vittate, the elytron slender antenna (with article 11 strongly acuminate), either avittate, faintly vittate, or distinctly vittate. front tarsal claws conspicuously asymmetrical, and The female cranium, femur, and the first five pygidium exceptionally robust and truncate will visible abdominal sterna are partly testaceous and distinguish males of this species from other males dark brown; sternum vi is always black. within the reventazon group. DISTRIBUTION (Figure 364).—The known range DESCRIPTION.—Form: As in P. reventazon (Figure extends from Mexico to Costa Rica with locality 186) except more slender. records from Guatemala and Honduras. Size: Length: male, 8.0 mm. Width: male, 2.4 mm. LOCALITY RECORDS (Figure 364).—I examined 98 adult speci- Color: As in males of P. calcaris except coxae not mens from Central America. MEXICO: Estado de Nayarit: infuscated. Tepic (MNHB, 1 male); Estado de Mexico; Toluca (ZMAN, 1 female). Estado de Veracruz: Atoyac (BMNH, 1 female); 49 Head: As in P. reventazon except frons propor- miles [78 km] south of Acayucan (GEKI, 1 female); Orizaba tionally narrower (HW/IOW, 2.5), antenna more (BMNH, 1 male; CASC, 4 males and 4 females; GEKI, 3 slender, antennal article 11 more acuminate, and NUMBER 227 79 length/width ratio of each antennal article 2.0: DESCRIPTION.—Form: As in P. reventazon (Figure 1.3:1.7:1.7:1.7:1.9:1.9:1.9:1.6:1.4:2.1. 186). Pronotum: As in P. reventazon except less trans- Size: Length: male, 8.0 mm; females, average verse (PW/PLS, 0.91). about 8.5 mm, range 7.9-9.2 mm. Width: Male, Elytron: As in P. reventazon. 2.2 mm; females, average about 2.4 mm, range 2.0- Abdomen: Male pygidium conspicuously robust 2.8 mm. One male and 2 females measured. and truncate. Color: As in P. calcaris. Front Tarsal Claws: Boldly asymmetrical. Head: As in P. reventazon except antenna pro- Male Genitalia (Figures 235, 256): Strongly pig- portionally shorter (AL/PLS, 1.7). mented. Combined length of phallobase and Pronotum: As in P. reventazon except disc more phallobasic apodeme 6.3 times longer than para- convex and side margin of pronotum proper less mere. Combined length of paramere and pallobase arcuate. 3 times longer than phallobasic epodeme. Para- Elytron: As in P. reventazon. mere: outer margin compressed at middle; feebly Abdomen: Male pygidium subquadrate; pygidial curvate in apical fourth in lateral view; lateral apodeme as long as pygidial plate. depressions well depressed; dorsum convex and Front Tarsal Claws: Male, boldly asymmetrical; venter concave at their respective apical half, basal female, feebly asymmetrical. half dorsum and venter plane; mesoventral and Male Genitalia (Figures 237, 238): Strongly pig- mesodorsal margin sinuous. Sinus: dorsal narrow in mented. Combined length of phallobase and phallo- apical half, broad at basal half; ventral narrow, basic apodeme 4.3 times longer than paramere. Combined length of paramere and phallobase 5 constricted medially. Phallus: as in P. claudus times longer than phallobasic apodeme. Paramere: except more slender. conspicuously broad in basal four-fifths, abruptly NATURAL HISTORY.—W. R. M. Mason collected narrow apical fifth, lateral depression inconspic- the only available specimen of this species in June, uous in lateral view; dorsum broadly convex, in Chiapas, Mexico, at an altitude of 914 meters. explanate medially; venter broadly concave; meso- ETYMOLOGY.—Latin, the adjective exilis (thin). I ventral margin concave; mesodorsal margin feebly refer to the slender form of the antenna. sinuous and briefly denticulated preapically. Sinus: LOCALITY RECORDS (Figure 363).—I examined 1 adult speci- dorsal narrow in apical half, slightly expanded in men from Central America. MEXICO: Estado de Chiapas: studied (1 specimen examined). 20 miles [32 km] north of Huixtla (CNCC, 1 male, holotype). Female Genitalia: As in P. nigriventris. Internal Reproductive Organs: Female, as in 29. Perilypus latilobus, new species Figure 55; ovary composed of 12 ovarioles (2 speci- basal half; ventral subelliptical. Phallus: not FIGURES 237, 238, 363 mens examined). VARIATION.—Structural: The female specimen TYPE-LOCALITY.—Six kilometers northwest of from Tuxtla Gutierrez is more robust than the one Pueblo Nuevo, Rio Bahada, Estado de Chiapas, from Pueblo Nuevo. Mexico. Color: The specimen from Tuxtla Gutierrez has TYPE-SPECIMENS.—The holotype male is depos- a short narrow vitta on the elytral, disc which is not ited in NMNH, the allotype in CNCC. The first the case in the other two specimens studied. of these specimens was collected by G. H. Nelson in NATURAL HISTORY.—In June in Tuxtla Gutierrez, 1965, the second by J. M. Campbell in 1969. One Mexico, at 1280 meters, I collected one adult speci- paratype: GEKI, 1 female. men by beating roadside vegetation. G. H. Nelson DIAGNOSIS.—The combination of femora pre- collected the holotype in July, by sweeping road- dominantly black to dark brown and pronotal side vegetation near Rio Bahada, in Pueblo Nuevo, discal vitta abruptly constricted at subapical depres- Mexico. sion will generally distinguish members of this DISTRIBUTION (Figure 363).—Known only from species from the very similar specimens of P. nigri- southern Mexico. ventris, P. calcaris, and P. exilis. ETYMOLOGY.—The trivial name is a Latin com- 80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

pound noun formed from the adjective latus (wide) vex, subapical depression more sinuous, and side + the noun lobus (lobe). I refer to the lobe-like margins less arcuate. appearance of the paramere. Elytron: As in P. reventazon except elytron proportionally shorter (EL/EW, average about 4.4, LOCALITY RECORDS (Figure 363).—I examined 3 adult specimens from Central America. MEXICO: Estado de Chiapas: range 4.3-5.6; 20 specimens measured). four miles [6 km] northwest of Pueblo Nuevo, Rio Bahada Front Tarsal Claws: Symmetrical in both sexes. (NMNH, 1 male, holotype); Tuxtla Gutierrez (GEKI, 1 fe- Male Genitalia (Figures 240-242): Strongly pig- male): junction between routes 190 and 195 (CNCC, 1 female). mented. Combined length of phallobase and phallobasic apodeme variable in relation to length of paramere (see "Variation"). Combined length of 30. Perilypus distinct™ (Chevrolat), paramere and phallobase 3.0 times longer than new combination phallobasic apodeme. Paramere: apex boldly FIGURES 239-242, 363 uncinate; lateral depression broad and deeply impressed; dorsum convex, strongly braced at base; Derestenus distinctus Chevrolat, 1874:289 [lectotype: female, venter narrow, concave; mesoventral and mesodorsal deposited in MNHP, here designated; type-locality: Yuca- margins sinuous. Sinus: dorsal broadly lanceolate; tan, Estado de Yucatan, Mexico].—Gorham, 1882:143.— Schenkling, 1906:262. ventral elliptical, constricted basally. Phallus: abruptly narrowed near apex; marginal denticles DIAGNOSIS.—Within the reventazon group speci- prominent (18 specimens examined). mens from between the southern half of Mexico Female Genitalia: As in P. nigriventris. and north of Honduras belong to this species if Internal Reproductive Organs: Male, anterior they are particularly densely pubescent, have the accessory gland uniramous; coiled posterior acces- pronotal sides broadly flavotestaceous and often sory gland biramous, divided into inner and outer roseate, and have a short but robust body form. branch (3 specimens examined). DESCRIPTION.—Form: As in Figure 239. VARIATION.—Structural: The ratio between the Size: Length: males, average about 8.5 mm, range combined length of the phallobase and the phallo- 6.3-9.4 mm; females, average about 8.1 mm, range basic apodeme over the length of the paramere 6.5-9.8 mm. Width: males, average about 2.0 mm, is highly variable (6.5 to 9.0). The variation may range 1.8-2.6 mm; females, average about 2.5 mm, be geographic but the available data are inconclu- range 2.0-2.8 mm. Ten males and 10 females sive. In general, the aforementioned ratio is lower measured. in males from Oaxaca than in those from Veracruz. Color: Clypeus, frons, and venter of cranium Color: Sex dimorphism is evident in the colora- flavotestaceous; epicranium predominantly black or tion of the abdominal venter and in the elytron. In predominantly flavotestaceous, often marked by females, the abdominal venter is bicolorous and the subquadrate black macula; gena black or flavo- elytron vittate. In males, the abdominal venter is testaceous; antenna black; pronotum with black concolorous and the elytron avittate. Of 17 females discal vitta that narrows at middle; pronotal sides examined 2 have the elytral vitta feebly incurved broadly flavotestaceous or roseate; pronotal collar apically. black; prosternum flavotestaceous; meso- and NATURAL HISTORY.—Adult specimens were col- metasternum dark brown or black; femora lected in Mexico from May to August. H. F. predominantly flavotestaceous, infuscated dorso- Howden collected 9 specimens at 609 meters by apically; tibia and tarsus dark brown or black; beating tree foliage in Oaxaca, Mexico. elytral disc with or without narrow flavotestaceous DISTRIBUTION (Figure 363).—The known range of vitta that when present begins at midbase, gradu- this species extends from Veracruz, Mexico, to Alta ally oblique toward suture, and usually not in- Verapaz, Guatemala. curved apically; abdominal venter entirely black, or first five visible sterna flavotestaceous, sixth black. LOCALITY RECORDS (Figure 363).—I examined 45 adult speci- Head: As in P. reventazon except antenna more mens from Central America. MEXICO: Estado de Veracruz: Cordoba (BMNH, 1 female, CASC, 1 male; GEKI, 1 male and serrate and proportionally shorter (AL/PLS, 1.6). 1 female; FMNH, 1 female); Montepio, 8 miles [13 km] north Pronotum: As in P. reventazon except more con- of Sontecomapan (GEKI, 1 female); Lago de Catemaco NUMBER TZ1 81

FIGURES 239-245.—Perilypus distinctus: 239, habitus; 240, tegmen, lateral view; 241, tegmen, ventral view; 242, phallus. P. crassus: 243, habitus; 244, antenna. P. ordinatus: 245, male pygidium. (Scale = 1 mm) 82 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(CNCC, 5 males and 1 female; GEKI, 2 males and 2 females); rate, flattened, and profusely pubescent; scrape Cerro de Palma (BMNH, 1 female). Estado de Oaxaca: 6 shorter than article 11; and length/width ratio of miles [10 km] south of Valle Nacional (CNCC, 1 male; each male antennal article 2.0:1.0:1.0:1.1:1.1:1.0: GEKI, 4 males and 1 female; HFHO, 4 males and 1 female). Estado de Chiapas: 20 to 25 miles [32 to 40 km] north of 1.0:1.0:1.0:1.0:1.8. Huixtla (CNCC, 1 female); Rio de Salvador, 4 kilometers Pronotum: As in P. reventazon except only feebly northwest of Ixhautan (GHNE, 1 female). Estado de Yuca- transverse (PL/PW, average about 0.93, range 0.91- tan: Yucatan (MNHP, 1 female, lectotype). "Mexico" 0.95; 3 specimens measured), integumental setae (MNHP, 1 male and 3 females; MNHB. 1 female; ZMAN, 1 male). GUATEMALA: Departamento de Alta Verapaz: more numerous and longer, disc more convex, sub- Teleman (BMNH. 2 males). "Guatemala" (GEKI, 1 male). apical depression more sinuous, and side margins BELIZE: Distrito de Corozal: Rio Hondo (GEKI, 1 male). of pronotum proper more arcuate. Distrito de Belize: Belize (GEKI, 1 male). "M-tee Dist" Elytron: As in P. reventazon except with more (GEKI, 1 female; MCZC, 1 male and 1 female). depth, prominently convex and truncated, surface REMARKS.—A male specimen formed part of the more arenose and more densely pubescent, epipleu- syntype series of P. nigriventris. Other specimens ral fold explanate dorsally, EL/EW, average about were identified as P. quadrilineatus, P. orthopleu- 4.4, range 4.3-4.6 (3 specimens measured). ridus, P. nigrifrons, and Colyphus signaticollis Front Tarsal Claws: Feebly asymmetrical in both Spinola. sexes. Male Genitalia: Strongly pigmented. Combined 31. PerUypus crassus, new species length of phallobase and phallobasic apodeme 5.7 times longer than paramere. Combined length of FIGURES 243, 244, 246-248, 365 paramere and phallobase 3.4 times longer than phallobasic apodeme. Paramere: apex feebly up- TYPE-LOCALITY.—Rio Huallaga, Yurimaguas, curved; lateral depression particularly prominent; Chambireyacu, Provincia de Huallaga, Peru. dorsum convex; explanate medially; venter con- TYPE-SPECIMENS.—The holotype male and the cave; mesoventral margin arcuate; mesodorsal mar- allotype, deposited in MNHP, were collected by gin feebly arcuate in most of posterior two-thirds, M. de Mathan in 1885. One paratype: GEKI, 1 strongly arcuate in anterior third. Sinus: dorsal male. feebly elliptical at middle, oval basally; ventral el- DIAGNOSIS.—The combination of robust body that liptical. Phallus: apex feebly digitiform; marginal is densely pubescent and truncated, boldly serrate denticles prominent (2 specimens examined). antenna (Figure 244), and South American distribution will easily identify members of this species Female Genitalia: As in P. nigriventris. within the reventazon group. VARIATION.—Structural: The female antenna is DESCRIPTION.—Form: As in Figure 243. slightly more serrate than male antenna. Size: Length: males, 7.4 mm; female, 7.3 mm. Color: The pronotal discal vitta is narrow me- Width: males, 4.6 mm; female, 4.3 mm. Two males dially in the females and broad throughout in and 1 female measured. males. Color: Clypeus, anterior half of frons, and venter NATURAL HISTORY.—Unknown except M. de of cranium navotestaceous; posterior half of frons Mathan collected two specimens near Rio Huallaga, black; epicranium predominantly black, fulvous in Peru, one in June and one in August. postocular region; prosternum flavotestaceous; pro- DISTRIBUTION (Figure 365).—Known only from notum flavotestaceous dorsolaterally, black ventro- Huallaga to Loreto, Peru. laterally and discally; pronotal collar black; front ETYMOLOGY.—Latin, the adjective crassus (stout), femur flavotestaceous in proximal half, dark brown with reference to the short stout body of this in- in distal half; middle and hind femur flavotestace- sect. ous in basal two-thirds, dark brown in distal third; elytron piceous but epipleural fold slightly paler; LOCALITY RECORDS (Figure 365).—I examined 3 adult speci- abdominal venter black. mens from South America. PERU: Provincia de Huallaga: Chambireyacu, Yurimaguas, Rio Huallaga (MNHP, 1 male, Head: As in P. reventazon except antennal ridge holotype, and 1 female). Provincia de Loreto: Rio Amazonas more prominent; antenna (Figure 244) more ser- (GEKI, 1 male). NUMBER 227 83

32. Perilypus immitis, new species to extent that mesodorsal margins overlap; venter concave; mesoventral margin and mesodorsal mar- FICURES 249-253, 364 gin sinuous, latter denticulated in apical half only. TYPE-LOCALITY.—Atlixco, Estado de Puebla, Sinus: dorsal indistinct, ventral obovate and acumi- Mexico. nate basally. Phallus: phallic plate abruptly narrowed in apical fourth; marginal denticles localized TYPE-SPECIMENS.—The holotype male and allo- at middle third of plate (5 specimens examined). type are deposited in CASC. Five paratypes deposited in repositories as noted under "Locality Rec- Female Genitalia (Figures 252, 253): Dorsal and ords." ventral lamina short; proctiger particularly broad; DIAGNOSIS.—Within the reventazon group, speci- remainder as in P. nigriventris (1 specimen exam- mens from Mexico belong to this species if their ined). elytra are predominantly or entirely blue-green or VARIATION.—Structural: The epicranium and olive green (the elytral apex is usually testaceous), frons are more coarsely punctated in males than in and if their elytral surface is densely vested with females. gray reclinate setae. Color: The male specimen from Yecora does not DESCRIPTION.—Form: rectangulate; these beetles have the elytral apex testaceous. Also, in this beetle are exceptionally small and shallow-bodied. the elytron has a more bluish tinge; in most other Size: Length: males, average about 6.2 mm, range specimens, the elytron is olive green. 5.4-6.9 mm; females, average about 6.9 mm, range NATURAL HISTORY.—In May, in Yecora, Mexico, 5.9-8.0 mm. Width: males, average about 2.2 mm, H. F. Howden collected one adult specimen at range 1.6-3.0 mm; females, average about 2.3 mm, 2134 meters by beating oak foliage (Quercus sp.). range 2.0-2.6 mm. Five males and 2 females meas- Specimens were also collected in June and July. ured. ETYMOLOGY.—Latin, the adjective immitis Color: Cranium entirely testaceous, or clypeus (rough). The name refers to the coarse dentitions frons and venter of cranium testaceous and epi- along the mesodorsal margin of the parameres. cranium and gena black; antenna dark brown; DISTRIBUTION (Figure 364).—From Sonora to prothorax testaceous, except pronotum with mid- Puebla, in Mexico. apical dark brown macula; mesosternum predom- LOCALITY RECORDS (Figure 364).—I examined 7 adult speci- inantly testaceous; metasternum black; femora mens from Central America. MEXICO: Estado de Sonora: becoming more dark brown from front to middle Yecora (GEKI, 1 male). Estado de Queretaro: San Juan del Rio (GEKI, 1 female; MNHB, 1 male). Estado de Puebla: femur; elytron olive green or blue-green; abdomen Atlixco (CASC, 1 male, holotype, and 1 female; GEKI, 1 testaceous or dark brown. male). Estado de Morelos: Tepoztlan (MHNM, 1 male). Head: As in P. reventazon except eyes less convex and cranium more coarsely punctated (particularly in males). 33. Perilypus caliculus, new species

Pronotum: As in P. reventazon. FIGURES 254-256, 363 Elytron: As in P. reventazon except elytron proportionally shorter (EL/EW, average about 4.2, TYPE-LOCALITY.—Tejupilco, Estado de Mexico, range 3.2-4.7; 11 specimens measured) and more Mexico. densely vested with reclinate setae. TYPE-SPECIMENS.—The holotype male, deposited Front Tarsal Claws: Male, boldly asymmetrical; in FMNH, and 73 paratypes were collected by female, feebly symmetrical. H. E. Hinton and R. L. Usinger in 1933. Paratypes Male Genitalia (Figures 249-251): Strongly pig- are deposited in repositories as noted under "Lo- mented. Combined length of phallobase and phallo- cality Records." basic apodeme 5 times longer than paramere. Com- DIAGNOSIS.—The male genitalia (Figures 254- bined length of paramere and phallobase 2.7 times 256) provides the only reliable characteristics for longer than phallobasic apodeme. Paramere: very diagnosing members of this species. broad basally; lateral depression deeply impressed DESCRIPTION.—Form: As in P. reventazon (Figure at middle third; dorsum convex, explanate medially 186). SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 84

FIGURES 246-259.—Perilypus crassus: 246, tegmen, lateral view; 247, tegmen, ventral view; 248, phallus. P. immitis: 249, tegmen, lateral view; 250, tegmen, ventral view; 251, phallus; 252, ovipositor, ventral view; 253, ovipositor, dorsal view. P. caliculus: 254, tegmen, lateral view; 255, tegmen, ventral view; 256, phallus. P. carbonarius: 257, tegmen, lateral view; 258, tegmen, ventral view; 259, phallus. (Scale = 1 mm) NUMBER 227 85 Size: Length: males, average about 7.8 mm, range Color: Among the males studied 60 have the 5.9-9.0 mm; females, average about 7.6 mm, range cranium entirely flavotestaceous and the prothorax ti.4-9.2 mm. Width: males, average about 2.3 mm, predominantly flavotestaceous; in other specimens range 2.1-2.6 mm; females, average about 2.5 mm, the cranium and prothorax is predominantly black. range 2.3-3.0 mm. Ten males and 10 females meas- In females, the cranium is always mostly black; the ured. pronotum either predominantly flavotestaceous and Color: Cranium entirely flavotestaceous, or clyp- maculated midapically or predominantly black, in eus postocular region of epicranium and cranial which case it has a black discal vitta. venter flavotestaceous and frons and epicranium I assign 14 specimens from Tejupilco, Mexico, to black; prothorax flavotestaceous, with or without a trivittate phenon. These have the elytron trivit- piceous macula at pronotal midapex, or prosternum tate; the vitta on the epipleural margin extends and upper sides of pronotum and pronotal disc from the elytral base to the elytral apical half or black; mesoscutellum flavotestaceous or black; more; the discal vitta begins at the elytral base meso- and metasternum from flavotestaceous to (nearer the posthumeral margin than to the sutural black; legs predominantly flavotestaceous or pre- margin) curves inward at the basal fifth and grad- dominantly black; elytron usually piceous or black ually outcurves to the elytral apical fifth where it and avittate, rarely flavotestaceouso r vittate; vittate curves inwards towards the elytral apex; the su- elytron with flavotestaceous vitta on epipleural mar- tural vitta is faintly developed and extends from gin, disc, and sutural margin. the elytral base to the elytral basal third. Head: As in P. reventazon except setiferous punc- NATURAL HISTORY.—Adult specimens were col- tations on frons and epicranium larger and antenna lected from Mexico in May; in June by H. E. more serrate. Evans from Morelos at 1829 meters and by G. H. Pronotum: As in P. reventazon except subapical Nelson on Mimosa sp. in Valle de Bravo; in July depression more deeply impressed. by H. E. Hinton from Tejupilco at 1164 meters; Elytron: As in P. reventazon except elytral sur- in August by G. H. Dieke from Michoacdn at 1378 face more arenose. meters; and in October by H. H. Smith from Chil- Front Tarsal Claws: Symmetrical in both sexes. pancingo at 378 and 914 meters. Male Genitalia (Figures 254-256): Strongly pig- DISTRIBUTION (Figure 363).—A specimen from mented. Combined length of phallobase and phallo- Turrialba, Costa Rica, extends the known range of basic apodeme about 7 times longer than paramere. this species from Mexico to Costa Rica. Combined length of paramere and phallobase about ETYMOLOGY.—Latin, a masculine diminutive 3.0 times longer than phallobasic apodeme. Para- noun caliculus (cup). I refer to the cup-shaped mere: arcuate in lateral view; dorsum convex; ven- contour of the combined venter of the parameres. ter broadly concave; mesoventral and mesodorsal margin concave. Sinus: dorsal and ventral elliptical. LOCALITY RECORDS (Figure 363).—I examined 168 adult Phallus: phallic plates broad, with marginal den- specimens from Central America. MEXICO: Estado de Hidalgo: Metamoros (CASC, 2 males and 2 females; GEKI, 1 ticles faintly visible (74 males examined). male). Estado de Mexico: Tejupilco (FMNH, 36 males, holo- Female Genitalia: As in P. nigriventris except type, and 41 females; GEKI, 6 males and 6 females); Real de lateral lobes of ventral lamina proportionally Arriba (FMNH, 1 male and 4 females; GEKI, 1 male and 1 shorter. females); Valle de Bravo (GEKI, 3 males and 2 females; Internal Reproductive Organs: Male, anterior ac- GHNE, 10 males and 8 females); Cuernavaca (CASC, 2 males and 3 females; CNCC, 2 males and 1 female; CUNY, 1 male cessory gland uniramous, coiled; posterior accessory and 1 female; GEKI, 2 males and 2 females). Estado de gland biramous (5 specimens examined). Morelos: Yautepec (BMNH, 1 male and 1 female). Estado VARIATION.—Structural: The female pronotum is de Guerrero: Tepetlapa (BMNH, 1 male and 1 female; GEKI, usually more transverse than the male pronotum. 1 male); Chilpancingo (AMNH, 1 female; BMNH, 5 males The setiferous punctations on the head and pro- and 8 females; CNCC, 1 female; GEKI, 1 male; Iguala (CNCC, 1 male and 1 female); Soledad (BMNH, 1 male); notum are particularly robust in some specimens Tasco (BMNH, 1 female; NMNH, 1 male and 1 female). from Valle De Bravo, Mexico; also in some of "Mexico" (BMNH, 1 male). "Guatemala" (GEKI, 1 male). these specimens the interocular depressions are COSTA RICA: Provincia de Cartago: Turrialba (GEKI, 1 coarsely rugose. male). 86 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY REMARKS.—At first glance specimens of the tri- ocular region of epicranium flavotestaceous and vittate phenon are very different from the others remainder of cranium black; prothorax entirely included herein, but they do not differ appreciably flavotestaceous except for midapical piceous mac- in any structural characteristic studied. ula, or predominantly piceous and pronotal disc vittate; meso- and metasternum flavotestaceous, or from dark brown to black; legs flavotestaceous or 34. Perilypus carbonarius Spinola black, or both; elytron variable (see "Variation"); abdomen from dark brown to black. FIGURES 257-265, 36S Head: As in P. reventazon except antenna (Fig- Perilypus carbonarius Spinola, 1844a: 105 [holotype: male, ure 260) more serrate; epicranial setiferous punc- deposited in MSPI; type-locality: "Mexico"].—Ekis, 1975:26. tations larger; frons often rugose; and length/width Derestenus lateralis Chevrolat, 1874:289 [lectotype: female, ratio of each male antennal article 2.0:1.0:1.4:1.3: and 1 female paralectotype, here designated, deposited in BMNH; type-locality: "Mexico"; new synonymy].— 1.3:1.2:1.2:1.1:1.1:1.1:1.9. Gorham, 1882:142. Pronotum: As in P. reventazon except more trans- Derestenus mutabilis Chevrolat, 1874:289 [lectotype: female, verse in females, subapical depression more deeply and S female paralectotypes, here designated, deposited in impressed, setiferous punctations larger, and side MNHP; type-locality: "Mexico"; new synonymy].— Gorham, 1882:142. margin of pronotum proper more boldly arcuate. Derestenus nigrifrons Chevrolat, 1874:289 [lectotype: male, Elytron: As in P. reventazon except surface more here designated, deposited in MNHP; type-locality: arenose. Oaxaca, Estado de Oaxaca, Mexico; new synonymy].— Front Tarsal Claws: Symmetrical in both sexes. Gorham, 1882:142.—Wolcott, 1927a:35. Male Genitalia (Figures 257-259): Strongly pig- Colyphus flammeus Gorham, 1878:162 [lectotype, female, here designated, deposited in BMNH; type-locality: mented. Combined length of phallobase and phal- "Mexico"; Gorham, 1882:142, placed this nominal species lobasic apodeme about 5 times longer than para- in synonymy with Derestenus mutabilis Chevrolat]. mere. Combined length of paramere and phallobasic Colyphus marginatus Gorham, 1878:162 [lectotype: female, apodeme about 3.6 times longer than phallobasic here designated, deposited in MNHP; type-locality: "Mex- apodeme. Paramere: feebly arcuate in lateral view; ico"; Gorham, 1882:142, placed this nominal species in dorsum feebly convex, boldly braced basally; venter synonymy with Derestenus lateralis Chevrolat]. concave; mesodorsal and mesoventral margin con- See "Remarks" for the justification of the new cave. Sinus: dorsal and ventral lanceolate; dorsal synonymies. slightly broader and longer than ventral. Phallus: DIAGNOSIS.—Although somewhat variable, genital as in Figure 259 (38 specimens examined). characteristics provide the most reliable clues for Female Genitalia: As in P. nigriventris. identifying male members of this species (Figures Internal Reproductive Organs: Male; anterior ac- 257-259). In most cases Mexican specimens belong cessory glands uniramous, coiled; posterior acces- to this species if they possess the following com- sory glands biramous, divided into an inner and bination of characteristics: body deep, pronotum outer branch (1 specimen examined). boldly transverse, elytral surface densely arenose, VARIATION.—Structural: There is considerable from rugose, and cranium and pronotum impressed variation in the degree of concavity of the para- with large setiferous punctations. Color patterns of meral mesodorsal margin, and in width of the the elytron are also diagnostic in some individuals paramere and parameral dorsobasal brace. The ely- (Figures 262-265), particularly females. tral apex is sometimes acuminate at the suture as DESCRIPTION.—Form: As in Figures 260, 261. is the case in all females from El Palmito. The Size: Length: males, average about 8.0 mm, range pronotum is generally more transverse in females 6.8-10.0 mm; females, average about 8.6 mm, range than in males. 6.8-10.3 mm. Width: males, average about 2.3 mm, Color: Elytra are mostly concolorous, usually range 2.0-3.2 mm; females, average about 2.7 mm, black, rarely testaceous. When bicolorous the ely- range 2.4-3.3 mm. Ten males and 10 females meas- tron may have one of three color patterns, corre- ured. sponding to the marginate phenon, the trivittate Color: Cranium entirely flavotestaceous or en- phenon, or the subfaciate phenon. tirely black, or clypeus, cranial venter, and post- In specimens of the marginate phenon the ely- NUMBER 227 87 tron is broadly vittate at the posthumeral margin; elytral apex. In males, the same vitta may extend the vitta extends from the humeral angle to the from the humeral angle to the elytral apex, ter-

261

262 FIGURES 260-265.—Perilypus carbonarius: 260, habitus, lateral view; 261, habitus, dorsal view; 262-265, elytras. (Scale = 1 mm) 88 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY minate at midelytron, or be faintly visible at the mutabilis Chevrolat or Colyphus flammeus Gorham, elytral middle half. In one female specimen the while some of the marginate phenon were de- posthumeral vitta gradually extends towards the scribed as Derestenus lateralis Chevrolat or Coly- elytral apex (Figure 265). I studied 5 males and 11 phus marginatus Gorham. I believe the specimens females of this phenon. representing these phena are conspecific primarily The elytral color pattern of trivittate specimens because their elytral color characteristics intergrade is as described for the trivittate phenon of P. calicu- to some extent as do characteristics of the male lus (see "Variation"). However, in specimens of genitalia. this species the discal vitta is less incurved at the As a group, specimens of P. carbonarius are elytral basal fifth and is closer to the posthumeral variable in color and in male genitalia, but the margin; in two male specimens the vittae are only variation is unequivalent to interspecific variations faintly visible at the elytral base. I studied 5 males within Perilypus. Perhaps P. carbonarius is under- and 2 females of this phenon. going more rapid differentiation than congeneric The elytra of subfasciate specimens are illus- species and perhaps the observed interpopulation trated in Figures 262 and 263. The nonpiceous por- variations are manifestations of restricted gene flow tion of the elytral disc ranges from stramineous to and possibly of incipient speciation. Alternatively, rufotestaceous. I studied 1 male and 7 females P. carbonarius, as defined herein, may consist of of this phenon. several biological species in which stable external NATURAL HISTORY.—Specimens were collected gaps have not yet materialized. from Mexico in June, July, and August. H. F. How- The nominal species placed in synonymy by me den collected 8 adult specimens, by beating, from (see above) were considered "good" species by other El Palmito at 1251 meters. J. Laue collected 1 authors, primarily on the basis of color charac- adult from Volcan Colima at 1000 meters. teristics. I found such characteristics unsuitable for DISTRIBUTION (Figure 363).—From Sinaloa to inferring reproductive isolation within Perilypus. Oaxaca, Mexico.

LOCALITY RECORDS (Figure 363).—I examined 91 adult The chaletoides Group specimens from Central America. MEXICO: Estado de Sonora: Arroyo del Juchujaqui, Alamos (GEKI, 1 male); The members of this group are characterized by Guirocoba (GEKI, 1 male). Estado de Sinaloa: 15 miles [24 their stout body; small, shallow, and profusely dis- km] west of El Palmito (CNCC, 4 males and 2 females; GEKI, 1 male and 2 females); about 38 miles [61 km] east tributed elytral punctations; very arenose elytral of Villa Union (CNCC, 1 female; GEKI, 1 male and 1 female). surface; light coloration (predominantly castaneous Estado de Durango: Ventanas (GEKI, 1 male). Estado de or stramineous); and by the unusually long male Nayarit: El Pich6n (CISC, 1 female; GEKI, 1 male); Tepic posterior accessory glands. (MNHB, 1 female). Estado de Jalisco: Guadalahara (CMPP, The two species that comprise the group occur 3 males and 1 female; GEKI, 1 male and 1 female). Estado de in Mexico from Durango to Chiapas. Colima: Volcan Colima (GEKI, 1 male). Estado de Morelos: Yautepec (MNHB, 1 male). Estado de Veracruz: Cerro de Palmas (MNHB, 1 female); Playa Vicente (BMNH, 1 male and 1 female). Estado de Mexico: Bejucos (FMNH, 1 male); 35. Perilypus chaletoides, new species Temascaltepec (BMNH, 1 female). Estado de Guerrero: FIGURES 266-271, 274, 277, 278, 359 Acapulco (WFBA, 1 female); 8 miles [13 km] north of Iguala (GEKI, 1 male); 5 miles [8 km] south and 2.5 miles TYPE-LOCALITY.—Twenty-four kilometers north [4 km] east of Chilpancingo (GEKI, 1 female). Estado de Oaxaca: Jugila (BMNH, 5 males and 7 females; GEKI, 3 of LaVentosa, at junction with highway 185, Estado males and 3 females; MNHP, 5 females; MNHB, 1 male and de Oaxaca, Mexico. 4 females); Panixtlahuaca (BMNH, 3 males and 2 females); TYPE-SPECIMENS.—The holotype male is depos- GEKI, 3 males); Oaxaca (MNHP, 2 males and 1 female). ited in NMNH, the allotype in GHNE. Both speci- "Mexico" (BMNH, 2 females; GEKI, 4 males and 3 females; mens were collected by G. H. Nelson in 1965. MNHP, 3 males and 5 females; MNHB, 1 female; MSPI, 1 male, holotype). DIAGNOSIS.—The color pattern of the elytron as described and figured (Figure 266), will distinguish REMARKS.—Some specimens, here assigned to the members of this species from congeneric species. subfasciate phenon, were described as Derestenus DESCRIPTION.—Form (Figure 266): The specimens NUMBER 227 89 are broad and deep bodied, and conspicuously con- fourth abruptly narrowed in dorsal or ventral view, vex in lateral view. feebly declinate in lateral view; lateral depression Size: Length: males, average about 8.6 mm, range feebly impressed; dorsum plane, with narrow brace 7.6-9.9 mm; females, average about 9.7 mm, range at basal third (Figure 271); venter feebly concave; 8.0-10.6 mm. Width: males, average about 2.7 mm, mesoventral margin faintly arcuate; mesodorsal range 2.4-3.2 mm; females, average about 3.2 mm, margin sinuous in basal half, linear in apical half. range 2.6-3.6 mm. Nine males and 6 females meas- Sinus: dorsal and ventral lanceolate. Phallus: acum- ured. inate; marginal denticles on middle half of phallic Color: Cranium castaneous, with or without mid- plate (12 specimens examined). epicranial black macula; antenna black; prothorax Female Genitalia (Figures 267, 268): Dorsal lam- castaneous, except lower sides of pronotum and pro- ina bilobed, incision short; ventral lamina trilobed, notal discal vitta black; mesoscutellum castaneous, lateral lobes shorter than medial lobe; dorsal bac- black, or castaneous but infuscated distally; meso- culi not fused; ventral bacculus acuminate near and metasternum entirely black, or middle region middle; coxital plates absent (2 specimens exam- castaneous and lateral regions black; femur varia- ined). ble (see "Variation"); tibia and tarsus black; ely- Internal Reproductive Organs: Male (Figures tron narrowly black along sutural margin, black 277, 278); anterior accessory glands uniramous, at apical fifth, castaneous in remainder; abdomen coiled throughout; posterior accessory glands bi- black. ramous, outer branch about 1.5 longer than inner Head: Interocular depressions shallow; frontal branch, both branches exceptionally long and ex- umbo prominent; eyes moderately convex; HW/ tensively convoluted; testis composed of 12 folli- IOW, average about 2.3, range 2.0-2.3 (15 speci- cles. Female; spermathecal capsule bulbous distally mens measured); antenna (Figure 274) short; AL/ and spermathecal gland duct exceptionally long (2 PLS, 1.4, boldly serrate; length/width of each an- males and 2 females examined). tennal article sex dimorphic, male 2.7:1.4:1.6:1.5: VARIATION.—Structural: The front tarsal claws 1.2:1.2:1.2:1.0:0.88:0.72:1.6, female 1.8:1.2:1.3:1.1: are particularly asymmetrical in the male specimen 1.1:1.0:1.0:0.89:0.72:0.67:1.5. from El Camer6n, Oaxaca, Mexico. The interocu- Pronotum: Boldly convex and only slightly trans- lar depressions and disc of the pronotal arch are verse (PL/PW, average about 0.96, range 0.92-1.0; more coarsely punctated in male specimens than 15 specimens measured); subapical and prebasal de- in female specimens. Also correlated with sex is pressions deeply impressed; pronotal arch coarsely the degree of concavity of the epipleural margin punctate; side margins of pronotum proper boldly (best seen in the elytral lateral view), which is convex. conspicuously more concave in males than in fe- Mesoscutellum: Subquadrate. males. Elytron: Conspicuously convex and with excep- Color: The pronotal vitta may be percurrent, in- tional depth; epipleural margin straight in basal terrupted at the middle, or reduced to a midapical half, arcuate in apical half; punctations very small macula. In three male specimens the dark regions and profusely distributed; punctate and interpunc- of the elytron are confined to an elongate sutural tate surfaces conspicuously arenose; setal densely infuscation. The elytron of some specimens is pre- distributed; EL/EW, average about 4.2, range 3.3- dominantly flavocastaneous. 4.7 (15 specimens measured); slope of apical de- NATURAL HISTORY.—In August in Chiapas, Mex- flexion gradual. ico, C. W. O'Brien collected three specimens, by Front Tarsal Claws: Male and female feebly beating, at 885 meters. G. H. Nelson collected asymmetrical. specimens in July by sweeping roadside vegetation Male Genitalia (Figures 269-271): Strongly pig- in Oaxaca, Mexico. One specimen was collected mented. Combined length of phallobase and phallo- at light in July by Mastro and Schaffner. Other basic apodeme 5.4 times longer than paramere. Com- specimens were collected from Mexico in June. bined length of paramere and phallobase 3.0 times DISTRIBUTION (Figure 359).—Known only from longer than phallobasic apodeme. Paramere: apical southwestern Mexico. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 90

266

273

FIGURES 266-274.—Perilypus chaletoides: 266, habitus; 267, ovipositor, ventral view; 268, ovipositor, dorsal view; 269, tegmen, lateral view; 270, tegmen, ventral view; 271, tegmen, dorsal view; 274, antenna. P. bicolor: 272, tegmen, lateral view; 273, tegmen, ventral view. (Scale = 1 mm) NUMBER 227 91

ETYMOLOGY.—Latin compound adjective formed tio of each male antennal article 2.3:1.2:1.3:1.3:1.5: from Chaletus (a chrysomelid generic name) + the 1.3:1.1:1.0:1.0:1.0:1.5. suffix -oides (like). I refer to the similarity in the Pronotum: As in P. chaletoides. elytral color pattern between members of this spe- Elytron: As in P. chaletoides. cies and those of Chaletus. Front Tarsal Claws: As in P. chaletoides. LOCALITY RECORDS (Figure 359).—I examined 34 adult speci- Male Genitalia (Figures 272, 273): Strongly pig- mens from Central America. MEXICO: Estado de Oaxaca: mented. Combined length of phallobase and phal- 9 miles [15 km] east of El Cameron (TTUT, 1 male); 15 lobasic apodeme 5.7 times longer than paramere. miles [24 km] north of La Ventosa, at junction with highway Combined length of paramere and phallobase 3.2 185 (NMNH, 1 male, holotype; GEKI, 1 male and 1 female; times longer than phallobasic apodeme. Paramere: GHNE, 2 females); 8 miles [13 km] north of La Ventosa acuminate, abruptly narrowed at apical third; lat- (TAMT, 1 female); 14 miles [23 km] south of Matias Romero (GEKI, 1 female; TAMT, 2 females); Almoloya (NMNH, 1 eral depression narrow, deep, and projecting beyond male). Isthmus of Tehuantepec (MCZC, 1 female). Estado de basal limit of ventral sinus; dorsum plane; venter Chiapas: 4 miles [7 km] northwest of Pueblo Nuevo, Rio feebly convex; mesoventral margin concave; meso- Bajada (GHNE, 1 male); At junction between highways 190 dorsal margin feebly convex in basal third, linear in and 195 (HFHO, 1 male); Tuxtla Gutierrez (CNCC, 1 fe- apical two-thirds. Sinus: dorsal and ventral lanceo- male); Suchiapa (CISC, 1 male); Los Amates (AMNH, 1 late, dorsal slightly longer than ventral. Phallus: male); 21 miles [34 km] west of Lazaro Cardenas (CNCC, 1 lemale); 2.3 miles [4 km] west of Las Cruces (GEKI, 2 phallic plates broad in basal three-fourths, grad- males); 31 miles [50 km] southeast of Comitan (GEKI, 1 ually narrowed to apex in apical fourth; marginal male and 1 female; TAMT, 3 males and 1 female); 7 miles [11 denticles small, extended from basal fourth to km] southeast of Chiapa de Corzo (GEKI, 1 male); 19 miles [31 apical three-fourths of phallic plate (7 specimens km] west of Ocozocoautla (TAMT, 1 female); 7 miles [11 km] examined). southwest of Ocozocoautla (GEKI, 1 male and 1 female); 13 miles [21 km] northeast Cintalapa (GEKI, 2 males and 1 Female Genitalia: As in P. chaletoides. lemale); 2 miles [3 km] east Rizo de Oro (GEKI. 1 male). VARIATION.—Structural: No appreciable structural variation was noted. Color: The dorsal fascies is concolorous in the 36. Perilypus bicolor (Chevrolat), specimens studied, but femoral coloration is vari- new combination able and not correlated geographically or with re- FIGURES 272, 273, 275, 359 gard to sex. In four specimens, the front and middle femora are entirely testaceous, the hindfemur Sallea bicolor Chevrolat, 1874:287 [lectotype, female, deposited is predominantly testaceous and infuscated apically. in MNHP, here designated; type-locality: Veracruz, Estado In 6 specimens, the front and middle femur is de Veracruz, Mexico].—Gorham, 1882:144.—Schenkling, 1898:364. testaceous and infuscated apically, the hind femur entirely dark brown; in 14 specimens the front fe- DIAGNOSIS.—The dorsal fascies of these specimens mur is testaceous and infuscated apically, the mid- is entirely testaceous. dle and hind femur entirely dark brown. DESCRIPTION.—Form: As in P. chaletoides (Fig- Additional color variation involves the meso- ure 266). sternum which is lighter, as dark, or paler than the Size: Length: males, average about 8.2 mm, range metasternum. Also, in one female specimen the 7.1-8.9 mm; females, average about 8.6 mm, range epipleural fold is faintly infuscated at the mid- 7.3-10.2 mm. Width: males, average about 2.7 mm, elytron. range 2.4-3.0 mm; females, average about 2.8 mm, NATURAL HISTORY.—Specimens were collected in range 2.3-3.2 mm. Six males and 10 females southern Mexico from June through August. measured. DISTRIBUTION (Figure 359).—This Mexican species Color: Cranium, prothorax, and elytron testa- ranges from Durango to Oaxaca. ceous; antenna, meso- and metasternum, tibia, tarsus, and abdomen dark brown; femora variable (see LOCALITY RECORDS (Figure 359).—I examined 24 adult specimens from Central America. MEXICO: Estado de "Variation"). Veracruz: Veracruz (MNHP, 1 female, lectotype); Cotaxtla Head: As in P. chaletoides except antenna (Fig- Experiment Station, Cotaxtla (CISC, I female; GEKI, 1 male); ure 275) slightly less serrate, and length/width ra- Puente Nacional (CISC, 1 male); Motzorongo (MNHB, 1 92 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY female). Estado de Durango: Canelas (GEKI, 1 male). Estado Color: Clypeus, frons, and postocular region of de Guerrero: Taxco (GEKI, 1 female); Almolonga (GEKI, 1 epicranium flavotestaceous; remainder of cranium male). Estado de Oaxaca: Oaxaca (BMNH, 1 female; GEKI, dark brown; pronotal sides flavotestaceous, some- 1 male and 1 female; MNHP. 1 female). "Mexico" (FMNH, 1 times roseous; pronotal disc with piceous vitta; male and 1 female; GEKI, 1 male; MNHB, 1 male and 2 females; RNHN, 1 male; UZMD, 1 female; ZMAN, 2 females). thoracic sterna brown; legs variable (see "Varia- tion"); elytron predominantly dark brown, with marginal and discal flavotestaceous vittae; abdom- The quadrilineatus Group inal sternum variable (see "Variation"). The most striking characteristic of the species Head: Interocular depression broad, shallow, not included herein is their lampyroid body. Members crescentic; eyes boldly convex (HW/IOW, average of this group are also characterized by antenna about 2.2, range 2.0-2.4; 10 males and 10 females boldly serrate; elytron with arenose surface, small measured); antenna (Figure 276) serrate, not par- punctations, very gradual apical slope, and deflexed ticularly flattened, length/width ratio of each male posthumeral region deeply concave; and meso- antennal article 1.7:1.4:1.5:1.7:1.4:1.5:1.4:1.4:1.4: dorsal margin of paramere denticulated. 1.4:2.3; AL/PLS, 1.7. The quadrilineatus group exhibits more inter- Pronotum: Boldly transverse (PL/PW, average specific structural variation than the other species about 0.90, range 0.82-0.96; 20 specimens meas- groups do. Perilypus bicristatus, for example, has ured); sides of pronotum proper boldly arcuate; some particularly discordant characteristics includ- dorsum feebly convex; pronotal arch coarsely punc- ing, elytron bicarinate, mesoscutellum triangular, tate; subapical depression well impressed at sides, and pronotum concave medially and paralaterally. shallow at middle; pronotal foveae and prebasal The combined distribution of the five species depression deeply impressed; pronotal collar nor- included in this group extends from northeastern mal, not particularly narrow. Mexico to central Guatemala. However, only P. Mesoscutellum: Subquadrate. quadrilineatus extends north of the Isthmus of Elytron: Posthumeral margin arcuate in outline Tehuantepec where it occurs primarily along the and contour; narrow longitudinal region proximal Atlantic drainage systems. to epipleural fold gradually deflected, deflected region concave; punctations small, shallow, and evenly distributed throughout elytral disc although 37. Perilypus quadrilineatus (Chevrolat), diminished near suture; surface arenose and pro- new combination fusely vested with short reclinate setae; epipleural fold obliquely convex; epipleuron convex, explan- FIGURES 276, 279-283, 285, 286, 366 ate internally; apical slope very gradual; EL/EW, Derestenus quadrilineatus Chevrolat, 1843:14 [holotype: fe- average about 4.4 range 3.2-4.9 (20 specimens male, deposited in MNHP; type-locality: Veracruz, Estado measured). de Veracruz, Mexico].—Lacordaire, 1857:443.—Gorham, Front Tarsal Claws: Symmetrical in both sexes. 1882:141. Derestenus similis Thomson, 1860:57 [lectotype: male, here Male Genitalia (Figures 281-283): Strongly designated, deposited in MNHP; type-locality: Mexico]. sclerotized. Combined length of phallobase and phallobasic apodeme 5 times longer than paramere. DIAGNOSIS.—The bold, obliquely convex epipleu- Combined length of paramere and phallobase 2.3 ral fold and the narrow vitta of the elytral disc times longer than phallobasic apodeme. Paramere: (Figure 279) easily separate specimens of this species outer margin acutely inflected at apical third; apex from congeners within the quadrilineatus group. incurved; lateral depression elongate and deeply DESCRIPTION.—Form: As in Figure 279. impressed; dorsum convex, strongly explanate Size: Length: males, average about 10.1 mm, medially; venter narrow and concave; mesoventral range 9.0-10.9 mm; females, average about 10.6 margin concave; mesodorsal margin sinuous and mm, range 8.2-13.2 mm. Width: males, average denticulated at middle third. Sinus: dorsal narrow about 3.5 mm, range 3.0-4.4 mm; females, average in apical half, expanded in basal half; ventral el- about 3.5 mm, range 2.6-4.6 mm. Ten males and liptical. Phallus: phallic plates broad, feebly sclero- 10 females measured. tized subapically; marginal denticles extended from NUMBER 227 93

283

FIGURES 275-283.—Perilypus bicolor: 275, antenna. P. chaletoides: 277, posterior accessory gland; 278, male internal reproductive organs. P. quadrilineatus: 276, antenna; 279, habitus; 280, male internal reproductive organs; 281, tegmen, lateral view; 282, tegmen, ventral view; 283, phallus. (Scale = 1 mm) 94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY middle of phallic plate to phallic plicae (17 speci- studied were collected from the Atlantic side of the mens examined). Isthmus. Two specimens were collected from north- Female Genitalia (Figure 285, 286): Dorsal lamina ern Guatemala. bilobed; ventral lamina trilobed; proctiger short, LOCALITY RECORDS (Figure 366).—I examined 68 adult acute; proctigeral bacculus acuminate at posterior specimens from Central America. MEXICO: Estado de fifth; coxital plates absent (2 specimens examined). Tamaulipas: 5 miles [8 km] west of Gomez Farias (GEKI, 1 Internal Reproductive Organs: Male (Figure 280); female). Estado de San Luis Potosi: El Salto de Agua (GEKI, anterior accessory gland uniramous, coiled through- 1 female; 25 miles [40 km] north of Tamazunchale (CNCC, out; posterior accessory gland biramous, outer 2 females; GEKI, 2 females). Estado de Puebla: 15 miles [24 km] east of Teziutlan (CNCC, 1 male and 1 female; branch nearly twice as long as inner branch (1 GEKI, 1 female). Estado de Colima: Volcan Colima (GEKI, specimen studied). I female). Estado de Veracruz: Jalapa (GEKI, 1 female); VARIATION.—Structural: The angularity of the Fortin (FMNH, 2 males and 1 female); Los Amates (CNCC, 1 convex epipleural fold is more pronounced in some female); Tezonapa (FMNH, 1 female); Veracruz (BMNH, 1 specimens; in others it is the extent of its inner female; FMNH, 1 male; GEKI, 1 male; MNHP, 1 female, holotype); Cordoba (CASC, 3 males; GEKI, 1 male and 1 and outer explanation that is striking. In general, female; MNHP, 1 female); San Rafael, Jicaltepec (MCZC, 2 female antennae are more serrate than male anten- females; GEKI, 1 female); Cotaxtla Experiment Station, nae. Cotaxtla (CISC, 1 male and 1 female); Puente Nacional The geographically disjunct female specimen (GEKI, 1 female); Lago de Catemaco (CISC, 1 female; NMNH, from Volcan Colima, Mexico, differs from all other 1 female); 2 miles [3 km] north of Santiago Tuxtla (GEKI, 1 female); Cerro de Palmas (BMNH, 2 females; GEKI, 2 specimens studied as follows: pronotum more trans- females); Santiago Tuxtla (BMNH, 2 females); Playa Vicente verse (PL/PW, 0.81), with subapical depression (BMNH, 2 males); Cosamaloapan (BMNH, 1 female); Atoyac more deeply impressed, and with arch narrower and (MNHB, 1 female). Estado de Oaxaca: Oaxaca (GEKI, 1 more convex; elytral form more rectangulate elytral female); Temescal (GEKI, 1 female; GHNE, 2 males). Estado disc avittate; and legs entirely dark brown. Whether de Chiapas: Portugal, 7 miles [11 km] southeast of Simojovel these differences can be inferred to represent ge- (GEKI, 1 female); 8 miles [13 km] south of Simojovel (GEKI, 1 female); Santo Domingo, 15 miles [24 km] southeast of netic discontinuity or geographic variation will de- Simojovel (CISC, 1 female); Simojovel (CISC, 1 female); 10 pend on study of additional material from Colima miles [16 km] south of Malpaso (GEKI, 1 female). "Mexico" and its environs. Should this specimen (which I (BMNH, 1 female; GEKI, 3 females; MNHP, 2 males and labeled P. quadrilineatus (Chevrolat)) be judged 1 female; MNHB, 4 females; MZST, 1 female; NMNH, 2 distinct from P. quadrilineatus it would undoubt- females; ZMAN, 1 female). GUATEMALA: Departamento edly be a member of the sister of that species. de Alta Verapaz: Teleman (BMNH, I female). Color: The flavous femoral annulus varies in width, its definition is obliterated when the femur 38. Perilypus telephoroides (Gorham), is predominantly flavous. Coloration of the abdom- new combination inal venter varies with sex and is dark brown in males. In females, visible sterna I, n, and vi are FIGURES 284, 287-291, 366 brown, sterna in to v are flavotestaceous. Sometimes Colyphtis telephoroides Gorham, 1882:142 [lectotype; male, the female sternum m is partially flavotestaceous and two paralectotypes, male and female, here designated, and partially brown. deposited in BMNH; type-locality: Capetillo, Departamento de Sacatepequez, Guatemala].—Gorham, 1886:335.—Schenk- NATURAL HISTORY.—Specimens were collected ling, 1906:262. in Mexico in June by G. H. Nelson (by beating roadside vegetation in Oaxaca) and by H. S. Dybas DIAGNOSIS.—The short testaceous vitta on the from Veracruz at 920 meters. A female specimen elytral humeral angle (Figure 287) identifies speci- was intercepted in New Orleans (U.S.A.) on banana mens of this species. debris apparently transported from Mexico. Other DESCRIPTION.—Form: As in Figure 287. adult specimens were collected in Mexico in May, Size: Length: males, average about 8.6 mm, range June, and August. 6.2-10.4 mm; females, average about 9.4 mm, range DISTRIBUTION (Figure 366).—The center of dis- 7.9-10.1 mm. Width: males, average about 2.9 mm, tribution of this Mexican species appears to be the range 1.9-3.6 mm; females average about 3.2, range Isthmus of Tehuantepec: most of the specimens 2.6-3.6 mm. Eight males and 9 females measured. NUMBER 227 95

FIGURES 284-291.—Perilypus telephoroides: 284, antenna; 287, habitus; 288, ovipositor, ventral view; 289, ovipositor, dorsal view; 290, tegmen, lateral view; 291, tegmen, ventral view. P. quadrilineatus: 285, ovipositor, ventral view, 286, ovipositor, dorsal view. (Scale = 1 mm) 96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Color: Clypeus and postocular region of epicran- apically. Phallus: as in P. quadrilineatus (10 ium testaceous, remainder of cranium black; an- specimens examined). tenna black; pronotum testaceous dorsolaterally, Female Genitalia (Figures 288, 289): As in P. black ventrolaterally and discally; pro-, meso-, and quadrilineatus except dorsal lamina and its incision metasternum black; legs black; elytron black except broader and proctiger more robust (1 specimen with short testaceous vitta at basal third of post- examined). humeral margin (Figure 287); abdominal venter Internal Reproductive Organs: Male, anterior color variable (see "Variation"). accessory gland uniramous, coiled throughout. Head: Interocular depressions broad and well VARIATION.—Structural: Notable structural var- impressed; interocular surface coarsely rugose; epi- iation is evident in the length and degree of cranium densely punctate; eyes boldly convex; antennal serration. Male antennae are proportion- HW/1OW, average about 2.4, range 2.1-2.7 (20 ally longer and less serrate than female antennae. specimens measured); antenna (Figure 284) con- Color: The size of the postocular epicranial spicuously serrate and sex dimorphic with respect macula varies as does the length of the testaceous to length (AL/PLS, male 2.0, female 2.3); length/ vitta on the elytral humeral angle; in one specimen width ratio of each male antennal article 2.5:1.0: the vitta is broadened and extends from the humeral 1.1:1.4:1.2:1.1:1.1:1.1:1.0:1.0:1.8. angle to the elytral apex. Coloration of abdominal Pronotum: Distinctly transverse and feebly trape- venter is sex dimorphic being black in males; in zoidal; subapical depression shallow; pronotal arch females, visible sterna i-m are black, sterna IV-VI feebly convex and coarsely punctate; disc of testaceous. pronotum proper transversely wrinkled; PL/PW, NATURAL HISTORY.—In October in the interior average about 0.83, range 0.79-0.90 (17 specimens valley of Sierra de las Minas, Guatemala, R. D. measured). Mitchell collected one adult specimen at 1550 Elytron: Posthumeral margin straight in basal meters. Other adult specimens were collected from two-thirds, arcuate in apical third; narrow longi- Mexico in May and June. tudinal region proximal to epipleural fold acutely DISTRIBUTION (Figure 366).—The known range deflexed; concavity of deflexed posthumeral region extends from southern Mexico to southern elongate and deeply impressed; apical slope very Guatemala. gradual; surface copiously impressed with small LOCALITY RECORDS (Figure 366).—I examined 20 adult shallow punctations; punctate and interpunctate specimens from Central America. MEXICO: Estado de surfaces arenose and profusely vested with short Chiapas: Chilcultic, near El Rinc6n (CNCC, 1 male; GEKI, reclinate setae; epipleural fold unusually narrow; 1 male); Lagos des Calores, route 17 (HFHO, 1 male); El EL/EW, average about 4.1, range 3.9-4.9 (17 speci- Rincrtn, route 7 (GEKI, 2 females); 17 miles [27 km] south- mens measured). east of Teopisca, route 24 (CNCC, 1 female). "Mexico" (MNHP, 1 male, ZMAN, 1 female). GUATEMALA: De- Front Tar sal Claws: Male, feebly asymmetrical; partamento de Sacatepequez: Capetillo (BMNH, 1 male, female, symmetrical. lectotype, and 2 females; GEKI, 1 male and 1 female; MNHP, Male Genitalia (Figures 290, 291): Strongly pig- 1 female). Departamento de Zacapa: Santa Clara, north of mented. Combined length of phallobase and Cabanas (FMNH. 1 female). phallobasic apodeme 5 times longer than paramere. Combined length of paramere and phallobase 3.4 times longer than phallobasic apodeme. Paramere: 39. Perilypus coroniformis, new species apex obtuse; outer margin depressed at middle; FICURES 292-296. S66 lateral depression deep, extended from paramere base to paramere apical fourth; dorsum convex, TYPE-LOCALITY.—Eight kilometers west of San strongly explanate medially; venter narrow and Cristobal de las Casas, Estado de Chiapas, Mexico. concave; mesoventral margin concave in basal two- TYPE-SPECIMENS.—The holotype male and the thirds, sinuous in apical third; mesodorsal margin allotype are deposited in HFHO. These and 4 sinuous, denticulated in apical third only, and with paratypes were collected by H. F. Howden in 1969. conspicuous brace at base. Sinus: dorsal narrowly Four paratypes: GEKI, 1 male and 1 female; lanceolate; ventral elliptical and feebly constricted HFHO, 2 females. NUMBER 227 97

FIGURES 292-297.—Perilypus coroniformis: 292, antenna; 293, habitus; 294, tegmen, lateral view; 295, tegmen, ventral view; 296, posterior accessory glands. P. fluctns: 297, habitus. (Scales = 1 mm) 98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DIAGNOSIS.—The configuration of the pronotal than the female antenna. In general, the antennal discal vitta, as illustrated (Figure 293) and described articles are more transverse in females. is unique in Perilypus. Color: There is no conspicuous color variation DESCRIPTION.—Form: As in Figure 293. among the 4 specimens examined. Size: Length: males, average about 8.1 mm, range NATURAL HISTORY.—During June near San 8.5-10.7 mm; females, 10.0 mm. Width: males, Crist6bal de las Casas, Mexico, H. F. Howden average about 3.8 mm, range 3.6-4.0 mm; females, collected 6 adult specimens by beating oak (Quercus 3.6 mm. Two males and 2 females measured. sp.) foliage. Color: Clypeus testaceous, remainder of cranium, DISTRIBUTION (Figure 366).—Known only from the and antenna black; prosternum and pronotal sides type-locality. roseous; pronotal disc broadly vittate, vitta with ETYMOLOGY.—Latin noun corona (crown) + oblong roseous macula at center; meso- and noun forma (shape), and adjectival ending -is. The metathorax, legs, elytron, and abdomen black, name refers to the peculiar shape of the pronotal except femora testaceous ventroapically. discal vitta. Head: As in P. teleph.oroid.es except HW/IOW, LOCALITY RECORDS (Figure 366).—I examined 6 adult speci- average about 2.4, range 2.3-2.5 (4 specimens mens from Central America. MEXICO: Estado de Chiapas: 5 measured); antenna (Figure 292) boldly serrate; miles [8 km] west of San Cristobal de las Casas (GEKI, 1 male AL/PLS, sex dimorphic, male 2.7, female 2.3; and and 1 female; HFHO. 1 male, holotype, and 3 females). length/width ratio of each antennal article sex dimorphic, male 2.3:1.4:1.2:1.4:1.4:1.3:1.4:1.4:1.4: 40. Perilypus fiuctus,ne w species 1.4:1.9, female 2.4:1.0:1.2:1.3:1.2:1.2:1.1:1.0:1.0:1.0: 1.5. FIGURES 297, 301-303, 366 Pronotum: As in P. telephoroides except more TYPE-LOCALITY.—Sinanja, Departamento de Baja transverse (PL/PW, average about 0.80, range Verapaz, Guatemala. 0.77-0.84; 4 specimens measured), more trapezoidal, TYPE-SPECIMENS.—The holotype male and allo- and disc smooth. type, deposited in BMNH, were collected by G. C. Elytron: As in P. telephoroides except post- Champion. humeral margin arcuate, disc undulated longi- DIAGNOSIS.—Distinguishable from P. coroni- tudinally, and EL/EW, average about 4.0, range formis, the only other species in the quadrilineatus 3.8-4.2 (4 specimens measured). group with elytral surface undulated longitudinally, Front Tarsal Claws: Symmetrical in both sexes. by the testaceous annulation of the femora and by Male Genitalia (Figures 294, 295): Strongly pig- the sagittate pronotal vitta. men ted. Combined length of phallobase and DESCRIPTION.—Form: As in Figure 297. phallobasic apodeme 5 times longer than paramere. Size: Length: male, 8.7 mm; female, 9.0 mm. Combined length of paramere and phallobase 3 Width: male, 3.2 mm; female 3.5 mm. One male and times longer than phallobasic apodeme. Paramere: 1 female measured. outer margin broadly arcuate; apex subacute; Color: As in P. telephoroides, except posthumeral lateral depression well impressed, extended from margin avittate, pronotal vitta strongly constricted parameral base to parameral apical two-thirds; at subapical depression, and femora annulated. dorsum convex, strongly explanate mesally; venter Head: As in P. telephoroides. concave; mesoventral margin concave; mesodorsal Pronotum: As in P. telephoroides, except not margin convex and finely denticulated at middle transversely wrinkled, margins of pronotum proper third. Sinus: dorsal indistinct; ventral elliptical. boldly arcuate, and fovae more spheroid. Phallus: as in P. telephoroides (2 males examined). Elytron: As in P. telephoroides except post- Female Genitalia: As in P. telephoroides. humeral margin arcuate from base to apex, surface Internal Reproductive Organs: Male; as in Figure distinctly undulated longitudinally, and EL/EW 296 (1 specimen examined). 3.9 (2 specimens measured). VARIATION.—Structural: Antennal length varies Front Tarsal Claws: Symmetrical in both sexes. with sex; the male antenna is proportionally longer Male Genitalia (Figures 301-303): Strongly pig- NUMBER 227 99 mented. Combined length of phallobase and phal- ited in CNCC, was collected by H. F. Howden in lobasic apodeme 9 times longer than paramere. 1964. Combined length of paramere and phallobase 3.2 DIAGNOSIS.—Elytron bicristate (Figure 300), pro- times longer than phallobasic apodeme. Paramere: notum concave medially and paralaterally and apical region broadly uncinate in lateral view; with paramedial longitudinal elevations, mesoscutel- lateral depression shallow and localized at middle lum triangular, and frontal umbo particularly third; dorsum boldly explanate in apical half, feebly prominent (Figure 299). The frontal umbo is the explanate in basal half; venter narrowly concave; only region of body not black. mesoventral margin sinuous; mesodorsal margin DESCRIPTION.—Form: As in Figure 300. convex and coarsely denticulated in middle half. Size: Length 9.4 mm; width 2.3 mm. Sinus: dorsal indistinct; ventral broadly lanceolate. Color: Frontal umbo flavous, remainder of body Phallus: as in Figure 303. black. Female Genitalia: Not studied. Head: Cranium coarsely punctate; interocular VARIATION.—Structural: The two specimens are depressions deeply impressed and rugose posteriorly; structurally homogeneous. frontal umbo prominent, conspicuously convex, Color: The male abdominal venter is black; in and narrowly prolonged posteriorly; antenna female specimens visible sterna i-m are black and (Figure 298) boldly serrate and very flat, article 1 IV-VI testaceous. and 11 oblong, 2 to 9 transverse, 10 as long as DISTRIBUTION (Figure 366).—Known only from broad; length/width ratio of each antennal article Baja Verapaz, Guatemala. 1.2:0.8:0.9:0.8:0.7:0.8:0.9:0.9:0.8:1.0:1.9; AL/PLS, ETYMOLOGY.—Latin noun fluctus (wave), refer- 2.4. ring to the longitudinal undulation of the elytral Pronotum: Transverse (PL/PW, 0.83), coarsely disc. punctate; surface concave centrally and para- LOCALITY RECORDS (Figure 366).—I examined 2 adult speci- laterally, and with paramedial longitudinal eleva- mens from Central America. GUATEMALA: Departamento tions; subapical depression deeply impressed at de Baja Verapaz: Sinanja (BMNH, 1 male, holotype); San side margins, feebly impressed discally. Jeronimo (BMNH, 1 female). Mesoscutellum: Triangular. REMARKS.—The holotype was part of the syntype Elytron: Posthumeral margin arcuate; narrow series of P. telephoroides and can be easily distin- longitudinal region proximal to epipleural fold guished from members of that species by the more acutely deflexed; concavity of deflexed posthumeral convex margins of the pronotum and the elytron, region long and deeply impressed; apical slope very undulation of the elytral surface, testaceous annulus gradual; surface with small coarse punctations, of femur, and by the absence of an elytral humeral prominently arenose; disc bicarinate (Figure 300); vitta. epipleural fold conspicuously narrow; EL/EW, 3.7. The allotype was part of the syntype series of Front Tarsal Claws: Symmetrical. JP. ventralis. While there is considerable habitus NATURAL HISTORY.—The holotype was collected similarity between female members of the two from El Palmito, Mexico in July. species, orientation of the epipleural fold easily ETYMOLOGY.—Latin compound noun formed separates them. In specimens of P. ventralis the from the prefix bi- (two) + noun crista (ridge), epipleural fold is in the ventral position while in referring to the two conspicuous longitudinal specimens of the quadrilineatus group the fold is ridges on the elytral disc. lateral. LOCALITY RECORDS (Figure 366).—I examined 1 adult specimen from Central America. MEXICO: Estado de Sinaloa: 15 miles [24 km] west of El Palmito (CNCC, 1 female, holo- 41. Perilypus bicristatus, new species type).

FIGURES 298-300, 366 The ornaticollis Group TYPE-LOCALITY.—Twenty-four kilometers west of El Palmito, Estado de Sinaloa, Mexico. The species included herein are substantially TYPE-SPECIMENS.—The holotype female, depos- diverse anatomically, but as a group are charac- 100 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

299

302

FIGURES 298-303.—Perilypus bicristatus: 298, antenna; 299, head; 300, habitus. P. fluctus: 301, tegmen, lateral view; 302, tegmen; ventral view; 303, phallus. (Scale = 1 mm) NUMBER 227 101

FIGURES 304-307.—Perilypus ventralis: 304, antenna; 305, habitus; 306, elytron, X 16; 307, elytron, X 65 (arrow indicates microsculpture). (Scale = 1 mm) 102 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY terized by mesoscutellum transverse, body form (Figure 305); prosternum black; legs black except moderately to strongly oval, interocular depressions femur with testaceous annulus (Figure 56); meso- crescentic, clytral sutural notch right angle, elytral and metasternum black; elytron color variable (see punctations small, and elytral surface usually con- "Variation"); abdomen bicolorous, visible sterna spicuously arenose. Male genital characteristics i-m and vi black, iv-v flavous. include, combined length of paramere and phallo- Head: Gular process (Figure 308) filiform; inter- base less than twice as long as phallobasic apodeme, ocular depressions broad, shallow, conspicuously phallobase marked with two dorsal and two ventral rugose and only feebly crescentic; frontal umbo longitudinal bands, and first connecting membrane prominent, narrowed from clypeus to epicranium; vested with serrulated scales. antennal carina coarsely punctate; frons broad; Perilypus ventralis is a particularly aberrant eyes not particularly convex; HW/IOW, average member in the group. Only specimens of that about 2.2, range 2.0-2.3 (18 specimens measured); species have the mesoscutellum stalked, postscutellar antenna (Figure 304) serrate; length/width ratio of disc concave, epipleural fold in ventral position, antennal articles sex dimorphic, male 2.2:1.5:1.5: elytron undulated longitudinally, and anterior 1.7:1.3:1.2:1.2:1.2:1.2:1.0:1.9, female 2.3:1.5:1.7:2.0: accessory gland of male reproductive organs not 1.9:1.8:1.6:1.5:1.3:1.3:2.0; AL/PLS, 2.2. completely coiled. In general, there is more habitus Pronotum (Figures 305): Transverse (PL/PW, concordance between specimens of P. ventralis and average about 0.88, range 0.84-0.93; 18 specimens lampyriform members of the quadrilineatus group measured); densely setose; pronotal arch poorly than with the other members of the ornaticollis denned; subapical depression feebly impressed at group. The similarity, however, is due to conver- sides, nearly obliterated medially; pronotal foveae gence and is not an expression of immediate com- longitudinal and transversely rugose; prebasal mon ancestry. depression shallow; pronotal collar narrow. The composite distribution of the eight species Mesoscutellum (Figures 310, 312): Transverse, in this group extends from Ohio, U.S.A., to stalked, and concave discally. Distrito Puerto Caballo, Venezuela. Elytron: Shallow, undulated longitudinally and copiously vested with short setae; posthumeral margin arcuate; slope of apical deflection very 42. Perilypus ventralis (Gorham), new combination gradual; punctations small and profusely distributed FIC.URKS 304-319, 368 throughout surface (Figure 306); punctate and interpunctate surface arenose (Fig. 307); epipleural Colyphus ventralis Gorham, 1882:141 [lectotype: female, and fold broad, concave, and deflected internally; 3 paralectotypes, 2 males and 1 female, here designated, deposited in BMNH; type-locality: Duenas, Departamento epipleural fold (Fig. 311) as wide as antennal de Sacatepequez, Guatemala]. article 11; sutural notch right angle; apical slope gradual; EL/EW, average about 4.5, range 4.3-5.0 DIAGNOSIS.—In addition to the characteristics (18 specimens measured). given in the key, specimens of this species are easily Front Tarsal Claws: Symmetrical in both sexes. diagnosed by their large size (average length about Male Genitalia (Figures 313, 314): Feebly sclero- 13 mm), concave mesoscutellar disc (Figure 312), tized; phallobase with 2 dorsal and 2 ventral and by the narrow and longitudinal pronotal longitudinal bands, latter more sclerotized than foveae. remainder of phallobase. Combined length of DESCRIPTIONS.—Form: As in Figure 305. phallobase and phallobasic apodeme 8 times longer Size: Length: males, average about 12.9 mm, than paramere. Combined length of paramere and range 10.4-14.1 mm; females, average about 13.0 phallobase 1.4 times longer than phallobasic mm, range 10.0-14.4 mm. Width: males, average apodeme. Paramere: apex truncate; lateral depres- 4.6 mm, range 4.0-5.0 mm; females, average about sion absent; dorsum and venter plane; mesoventral 4.4 mm, range 3.6-5.2 mm. Eight males and 10 and mesodorsal margin broadly concave. Sinus: females measured. dorsal indistinct, half as long as lanceolate ventral. Color: Head including antenna black; pronotum First connecting membrane with serrulate scales flavous at sides, with black triangular discal vitta (Figure 315). Phallus: phallic plate very slender and NUMBER 227 10S

FIGURES 308-314.—Perilypus ventralts: 306, head venter and cervical membrane, X 90 (arrow indicates gular process); 309, elytron, X 41 (arrow indicates sutural cleft); 310, mesoscutellum, lateral view, X 45; 311, elytron, X 41; 312, mesoscutellum, dorsal view, X 45; 313, aedeagus, ventral view, X 60; 314, phallus. X 290. 104 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ficintzs 315-321.—Perilypus ventralis: 315, aedeagus, X 350 (arrow indicates first connecting membrane); 316, male internal reproductive organs; 317, ovipositor, ventral view; 318, ovipositor, dorsal view; 319, posterior accessory gland. P. ornaticottis: 320, aedeagus, ventral view, X 117; 321, aedeagus, ventral view, X 325 (arrow indicates first connecting membrane). (Scale - 1 mm) NUMBER 227 105 strongly sderotized; marginal denticles absent; specimens from Central America: MEXICO: Estado de Vera- phallic plicae prolonged and set with very large cruz: Fortfn (FMNH, 1 female). Jesus Carranza (NMNH, 1 spines (Figure 314) (18 specimens examined). female). Estado de Guerrero: Jalapa (BMNH, 2 males). Estado de Oaxaca: Oaxaca (BMNH, 1 male). "Mexico" Female Genitalia (Figures 317, 318): Dorsal lam- (MNHP, 1 female). GUATEMALA: Departamento de Baja ina heptalobed; ventral lamina trilobed, laminal Verapaz: San Jeronimo (GEKI, 1 female). Departamento de incisions very short; proctiger semicircular; proc- Sacatepequez: Antiqua (GEKI, 1 female; NMNH, 1 female); tigeral bacculi not fused; ventral bacculus acumi- Sumpango (AMNH, 1 female; GEKI, 5 males and 4 females; FMNH, 1 female; NMNH, 1 male and S females); Duenas nate near middle; coxital plates absent (2 specimens (BMNH, 4 females, lectotype; GEKI, 1 female). examined). Internal Reproductive Organs: Male (Figures 316, 319): anterior accessory gland uniramous, coiled 43. Perilypus ornaticollis (LeConte), distally, posterior accessory gland biramous, outer new combination branch slightly longer than inner branch, both branches broadly arcuate; ejaculary duct not bul- FIGURES 520-330, 367 bous anteriorly; testis composed of 12 follicles. Cleronomous ornaticollis LeConte, 1880:194 [holotype: female, Female: as in Figure 55 (4 males and 2 females deposited in MCZC; type-locality: Ohio, U.S.A.]. examined). Colyphus melanopterus Dury, 1906:251 [type deposited in VARIATION.—Structural: A female specimen from CMNH; type-locality: Cincinnati, Ohio, U.S.A.; Synonymy El Fortfn, Mexico, is more oval in body form. by Wolcott (1910a:851)]. Colyphus furcatus Schaeffer, 1904:218 [lectotype: male, and 7 Color: Intrapopulation variation of elytral color paralectotypes, 1 male and 6 females, here designated, de- not correlated with sex distinguishes a light phenon posited in NMNH; type-locality: San Tomas, Esperanza and a dark phenon. Beetles of the light phenon Ranch, Brownsville, Texas, U.S.A.; new synonymy]. have flavous vitta on the sutural and epipleural According to Dury (1906:251) the type-specimen margins. The epipleural vittae may be slightly of Colyphus melanopterus Dury (Cleronomus orna- expanded at midelytron. The elytron is entirely ticollis LeConte) is "longer, much less hairy, the black in beetles of the dark phenon. In general, the elytra are more shining and immaculate jet black" elytral disc varies from brown to black. than representatives of Colyphus furcatus Schaeffer. NATURAL HISTORY.—In June Henry Dybas col- Although I did not see the type-specimen of C. lected a female specimen from El Fortin, Mexico, melanopterus, two other specimens from Ohio were at 1000 meters. From Sumpango, Guatemala, at studied. The characteristics mentioned by Dury 2012 meters, I collected 15 adult specimens on the fall within the range of variation of this species. compositae Vernonia deppeana Lessing. Fourteen DIAGNOSIS.—Distinguishable by the bifurcate of these were of the dark phenon, one of the light configuration of the pronotal discal vitta (Figures phenon. On the same plant I also collected 322-324). specimens of the lampyrid genus Photinus. The DESCRIPTION.—Form: As in Figure 322. striking similarity between these beetles was first Size: Length: males, average about 6.7 mm, range noted by Gorham (1882:141), ". . . this, together 6.3-7.2 mm; females average about 7.9 mm, range with the color, gives this insect a singularly close 6.0-8.4 mm. Width: males, average about 2.0 mm, resemblance to some Lampyridae which the two range 1.6-2.4 mm; females, average about 2.3 mm, or three pale segments beneath heightens. The range 1.5-2.8 mm. Six males and 15 females variety from Oaxaca [my light phenon] equally measured. resembles those varieties of species of Photinus in Color: Clypeus and frons flavotestaceous; epi- which the margins are pale." cranium, gena, antenna, and legs dark brown; DISTRIBUTION (Figure 368).—The known distri- pronotum and elytron variable (see "Variation"); bution of this species extends from Veracruz, prosternum flavotestaceous; legs, meso- and meta- Mexico, to Sacatepequez, Guatemala. Most of the sternum, and abdomen brown to black. specimens were collected at elevations from 1000 Head: Interocular depressions broad, well im- to 2012 meters. pressed, and crescentic; frontal umbo prominent; LOCALITY RECORDS (Figure 368).—I have examined 28 adult frons broad; eyes moderately convex; HW/IOW, 106 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

325

FIGURES 322-330.—Perilypus ornaticollis: 322, habitus; 323-324, pronota; 325, antenna; 326, tegmen, lateral view; 327, tegmen, ventral view; 328, phallus; 329, ovipositor, ventral view; 330, ovipositor, dorsal view. (Scale = 1 mm) NUMBER 227 107 average about 1.8, range 1.3-1.9 (21 specimens also differ by their more transverse pronotum measured); antenna (Figure 325) boldly serrate; (PL/PW, 0.76). length/width ratio of antennal articles sex dimor- Color: In general, southern specimens are lighter phic, male 2.0:1.1:1.3:1.2:1.1:1.1:1.2:0.9:0.9:0.7:1.3, than northern ones. Beetles from the southern part female 2.0:1.1:1.3:1.2:1.2:1.2:1.0:1.0:0.8:0.7:1.2; AL/ of the range (south of Louisiana) have an incom- PLS, 1.7. plete pronotal vitta (Figure 322), and the elytron is Pronotum: Strongly transverse (PL/PW, average broadly vittate (Figure 322). The pronotal vitta is about 0.81, range 7.4-8.7; 21 specimens measured); percurrent and the elytral disc a vittate and more side margin of pronotum proper boldly convex; shiny in specimens north of Louisiana (Figure subapical depression parallel with anterior margin, 324). pronotal arch narrow, feebly convex; prebasal Two specimens, one from Tennessee and one depression shallow; pronotal collar narrow. from Arkansas, have the pronotal vitta prolonged Mesoscutellum: Transverse. unto the basal half of the pronotal arch (Figure Elytron: Posthumeral margin straight in basal 323). third, arcuate in apical two-thirds; slope of apical NATURAL HISTORY.—Adult specimens were collected in March, May, June, and July. R. Kirkton deflection gradual; punctations broad and shallow; collected one specimen in Arkansas in June within punctate and interpunctate surfaces with rugose a "cotton alfalfa strip." Dury (1906:251) collected microsculpture, latter makes elytral surface appear "one specimen while sweeping low vegetation in arenose; densely setose; EL/EW, average about 4.5, river bottom, July 5, 1905." range 4.1-5.6 (21 specimens measured). DISTRIBUTION (Figure 367).—This is the only Front Tarsal Claws: Symmetrical in both sexes. species of Perilypus prominently distributed north Male Genitalia (Figures 320, 321, 326-328): Fee- of Mexico. The known range extends from Ohio, bly sclerotized; phallobase with 2 dorsal and 2 ven- U.S.A., to Cordoba, Mexico. tral longitudinal bands, latter more sclerotized than remainder of phallobase. Combined length of LOCALITY RECORDS (Figure 367).—I examined 46 adult specimens from North and Central America. UNITED STATES: phallobase and phallobasic apodeme 7 times longer Ohio: Hamilton County (FMNH, 1 male); "Ohio" (MCZC. 1 than paramere. Combined length of paramere and female, holotype). Indiana: Tippecanoe County (GEKI, 1 fe- phallobase 1.4 times longer than pallobasic apo- male). Tennessee: Memphis (GEKI, 1 female). Louisiana: deme. Paramere: acuminate; inner walls of para- Alexandria (GEKI, 1 female). Arkansas: Mississippi County (WFBA, 1 female). Texas: Saint Thomas, Brownsville (GEKI, mere densely setose. Sinus: dorsal, half as long as 1 female); Brownsville, Esperanza Ranch (AMNH, 1 female; ventral. First connecting membrane set with serru- CASC, 1 female; BMNH, 1 female; FMNH, 3 females; GEKI, late scales (Figure 321). Phallus: apex lobiform; 2 males and 8 females; GHNE. 1 female; LMDR, 2 females; phallic plate abruptly narrowed at middle and at MCZC, 1 male and 1 female; SEMK, 2 females; NMNH, 2 males and 7 females; UZMD, 1 female). MEXICO: Estado junction with phallic apodeme; marginal denticles de Veracruz: San Rafael, Jicaltepec (GEKI, I female; MCZC, absent; phallic plicae inconspicuous (6 specimens 2 females); Cordoba (GEKI, 1 female); Orizaba (GEKI, 1 fe- examined). male); 17 miles [27 km] north of Nauxtla (GEKI, 1 female). Female Genitalia (Figures 329, 330): Dorsal and REMARKS.—The three female specimens discussed ventral lamina trilobed, latter with very short under "Variation" are probably not conspecific, incisions; proctiger broad, nearly semicircular; but I cannot decide on the basis of the avail- proctigeral bacculi not fused; ventral bacculus able information. I labeled these P. ornaticollis accuminate near middle; coxital plates absent (2 (LeConte)?. specimens examined). Internal Reproductive Organs: Male: lateral accessory gland uniramous, coiled throughout (1 44. Perilypus galbeus, new species specimen examined). FICURES 332, 337-339, 355, 368 VARIATION.—Structural: The antenna is more serrate and body form more oval in three female TYPE-LOCALITY.—Guatemala. specimens from southeastern Mexico; two of these TYPE-SPECIMENS.—The holotype male is depos- 108 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ited in MNHP and was collected by L. Conradt. specimens examined differ in shape of interocular The allotype is in BMNH. depression which is more crescentic in the male. DIAGNOSIS.—There is considerable habitus simi- Color: The two specimens do not vary apprecia- larity between specimens of this species and those bly in color. of P. prolixicornis and of P. pilatus. From the first DISTRIBUTION (Figure 368).—The known range of these, P. galbeus differs by its shorter antennal extends from Veracruz, Mexico, to Guatemala. article 11 (compare Figures 331, 332), and from P. ETYMOLOGY.—Latin noun galbeus (arm band), pilatus by the more oval and more convex elytron. in allusion to the testaceous annulus on the Also, the elytral surface of P. pilatus is vested with femora. exceptionally long vertical setae. LOCALITY RECORDS (Figure 368).—I examined 2 adult speci- DESCRIPTION.—Form: As in Figure 355. mens from Central America. MEXICO: Estado de Veracruz, Size: Length: male 7.5 mm, female 8.8 mm. Cordoba (BMNH, 1 female). GUATEMALA (MNHP, 1 male, Width: male 2.4 mm, female 3.4 mm. holotype). Color: Head black, except clypeus and mid- anterior region of frons flavotestaceous; antenna black; prothorax predominantly flavotestaceous, 45. Perilypus prolixicornis, new species pronotal disc with tapering black vitta; legs black FIGURES 331, 334-336, 368 except femur annulated distally; meso- and metathorax, elytron, and abdomen black. TYPE-LOCALITY.—Cordoba, Estado de Veracruz, Head: Interocular depressions and frontal umbo Mexico. as in P. ornaticollis; frons broad, eyes moderately TYPE-SPECIMENS.—The holotype male, deposited convex; antenna (Figure 332) boldly serrate; in CASC, was collected by A. Feyes. antennal length sex dimorphic (AL/PLS, male 2.5, DIAGNOSIS.—The only specimen studied can be female 2.1) and article 11 as long as preceding two; distinguished from congeneres by the length of length/width ratio of each antennal article, male antennal article 11 (Figure 331), which is as long 2.2:1.4:1.5:1.3:1.2:1.2:1.0:1.0:1.0:1.0:2.0. as the preceding three. Pronotum: Strongly transverse (PL/PW, average DESCRIPTION.—Form: As in P. galbeus (Figure about 0.77, range 0.76-0.78; 2 specimens measured); 355). subapical depression feebly sinuous, shallow at Size: Length, 7.5 mm; width, 2.4 mm. middle; pronotal arch narrow and shallow; foveae Color: As in P. galbeus. more transverse than punctiform; pronotum proper Head: As in P. galbeus except antennal article margin feebly arcuate; prebasal depression very 11 (Figure 331) proportionally longer; length/width shallow; pronotal collar very narrow medially. ratio of each antennal article, 2.0:1.1:1.0:1.3:1.3: Mesoscutellum: Transverse. 1.1:1.0:0.9:0.8:0.9:3.5; and AL/PLS, 4.9. Elytron: More convex and more arenose than Pronotum: As in P. galbeus except foveae in P. ornaticollis; posthumeral margin arcuate broader and shallower. throughout; slope of apical deflection gradual; Elytron: As in P. galbeus. epipleural fold feebly concave, as broad as antennal Front Tarsal Claws: Symmetrical. article 11; apex truncate; EL/EW, average about Mesoscutellum: Transverse. 4.2, range 3.8-4.5 (2 specimens measured). Male Genitalia (Figures 334-336): Generally as Front Tarsal Claws: Symmetrical in both sexes. in P. ornaticollis (LeConte) but differs from that Male Genitalia (Figures 337-339): As in P. species as follows: combined length of phallobase ornaticollis except paramere apex more gradually and phallobasic apodeme 10 times longer than acuminate; dorsal and ventral sinus very narrow, paramere; paramere apices excurved, ventral sinus latter sinus only slightly longer than dorsal sinus; only slightly longer and broader than dorsal sinus, and phallic plates uniformly slender. and phallic plate uniformly narrow. Female Genitalia: Not studied. ETYMOLOGY.—Latin adjective prolixus (stretched VARIATION.—Structural: Antennal length is sex out) + the noun cornu (horn) + 1 adjectival dimorphic; the male antenna is the longer. The two ending -is; referring to the long antennal article 11. NUMBER Til 109

337 338 339 340 341

t1 IGURES 331—1-341.—JTI . PerilypuM. c / HVLSIAOs proltxicomis: u i un.Mii// riu , 331JJ,I antenna, a«ii«_iuia; ,334 O-J~X,, tegmen I«.^IIILI, lateraI l view; 335, teginen, ventral vieww;; 336,, phallus.. P.P. galbeus: 332,, antenna; 337, , tegmenlegmen, , latera_..„.**l * view,*»„; . 338..,...., , tegmen^.^ ,,. ventral vieww; 339,, phallus.. PP.. floralis:floralis: 333,, antenna; 340,, legmentegmen,, lateral view; 341,, teementegmen., ventral view. (Scali_e ^= i1 mmmrn)\ SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 110

342

FIGURES 342-350.—Perilypus floralis: 342, habitus; 343, tegmen, lateral view; 344, tegmen, ventral view; 345, phallus; 346, ovipositor, ventral view; 347, ovipositor, dorsal view. P. pilatus: 348, ovipositor, ventral view; 349, ovipositor, dorsal view. P. antarius: 350, antenna. (Scale = 1 mm) NUMBER 227 111

LOCALITY RECORD (Figure 368).—I examined 1 adult speci- LOCALITY RKCORDS (Figure 368).—I examined 1 adult specimen from Central America. MEXICO: Estado de Veracruz: men from Central America. PANAMA: Provincia de Pan- Cordoba (CASC, 1 male, holotype). ama, Cerro Campana (NMNH, 1 female, holotype).

46. Perilypus pilatus, new species 47. Perilypus antarius, new species

FIGURES 348, 349, 351, 354, 368 FIGURES 350, 352, 353, 368 TYPE-LOCALITY.—Cerro Campana, Provincia de TYPE-LOCALITY.—San Pedro, Montes de Oca, Panama, Panama. Provincia de San Jose", Costa Rica. TYPE-SPECIMENS.—The holotype female, depos- TYPE-SPECIMEN.—The holotype female, deposited ited in NMNH, was collected by H. P. Stockwell in NMNH, was collected by C. H. Ballou in 1936. in 1970. DIAGNOSIS.—Elytral apex strongly upcurved DIAGNOSIS.—The stout black setae on the elytral (Figure 353). surface and the conspicuous divergence of sutural DESCRIPTION.—Form: As in Figure 352. margins at the elytral apical fifth are diagnostic Size: Length 7.8 mm. Width 2.5 mm. characteristics of this species. Color: As in P. ornaticollis except epipleural fold DESCRIPTION.—Form: As in Figure 354. flavotestaceous, pronotal vitta not furcate, elytral Size: Length: 8.4 mm. Width: 2.9 mm. disc avittate, femora annulated subapically, and Color: As in P. ornaticollis except pronotal vitta posterior margin of abdominal sterna flavotest- not furcate; pronotal sides roseate; apex and suture aceous. of elytron black; and femora with subapical Head: As in P. ornaticollis except antenna testaceous annulus. (Figure 350) proportionally longer (AL/PLS, 2.5); Head: As in P. ornaticollis except eyes more length/width ratio of each antennal article 2.2: convex (HW/IOW, 2.1); antenna (Figure 350) 1.1:1.2:1.9:1.7:1.5:1.5:1.3:1.3:1.1:1.9; and HW/ more compressed, more distinctly serrate, and IOW, 2.2. proportionally longer (AL/PLS, 2.0); and length/ Pronotum: As in P. ornaticollis except subapical width of each antennal article 2.0:1.0:1.1:1.3:1.2: depression more impressed and PL/PW, 0.73. 1.2:1.2:1.0:0.9:0.8:1.3. Mesoscutellum: Transverse. Pronotum: As in P. ornaticollis except subapical Elytron: As in P. ornaticollis except apex uplifted depression more sinuous and foveae more deeply epipleural fold obliquely convex; and EL/EW, 4.8. impressed. Front Tarsal Claws: Symmetrical. Mesoscutellum: Transverse. NATURAL HISTORY.—In June, from the type- Elytron: Posthumeral margin arcuate; puncta- locality, G. H. Ballou collected the holotype on tions particularly small and shallow; punctate and Hemalia erecta Linnaeus. interpunctate surfaces densely arenose; surface ETYMOLOGY.—Latin adjective antarius (hoisting), copiously vested with short pale reclinate setae and referring to the uplifted elytral apex. with stout black vertical setae; sutural hinge LOCALITY RECORDS (Figure 368).—I examined 1 adult speci- abruptly narrowed at apical fifth; elytral sutural men from Central America. COSTA RICA: Provincia de San margin diverging in apical fifth; apex sinuato- Jos£: San Pedro, Montes de Oca (NMNH, 1 female, holotype). truncate (Figure 354); EL/EW, 3.9. Front Tarsal Claws: Symmetrical. 48. Perilypus sinuapicis, new species Female Genitalia (Figures 348, 349): As in P. ornaticollis except dorsal laminal incisions shorter FIGURES 343-345, 369 and proctiger more slender. NATURAL HISTORY.—In July H. P. Stockwell TYPE-LOCALITY.—San Esteban, Estado de Cara- collected the holotype from the type-locality (800 bobo, Venezuela. meters) by beating. TYPE-SPECIMENS.—The male holotype, deposited ETYMOLOGY.—Latin adjective pilatus (thick), in FMNH, was collected by P. J. Anduze in 1939. referring to the stout elytral vertical setae. DIAGNOSIS.—Easily distinguishable from speci- 112 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

354

FIGURES 351-354.—Perilypus pilatus: 351, antenna; 354, habitus. P. antarius: 352, habitus; 353, hind body. (Scale = 1 mm) NUMBER 227 113 mens of P. fioralis, the only other member of the Color: As in P. ornaticollis except pronotal vitta ornaticollis group whose specimens have the elytral not furcate and femur with testaceous annulus. surface very shiny, by differences in elytral color Head: As in P. ornaticollis except eyes more con- (elytron is avittate in specimens of P. sinuapicis). vex (HW/IOW, 2.2), antenna proportionally longer Also, in specimens of P. sinuapicis the antenna is (AL/PLS, 2.1), and length/width ratio of each black and femora predominantly dark brown antennal article 2.0:1.1:1.4:1.5:1.5:1.3:1.2:1.2:1.1: whereas in those of P. fioralis the antenna and 1.2:1.5. the abdomen is entirely flavotestaceous. Pronotum: As in P. ornaticollis except less trans- DESCRIPTION.—Form: As in P. fioralis except verse and subapical depression more deeply epipleural margin less convex. impressed. Size: Length: 6.2 mm. Width: 1.9 mm. Mesoscutellum: Transverse. Elytron: As in P. ornaticollis except punctate and interpunctate surfaces shiny, only feebly arenose, epipleural fold convex, and apex sinuatotruncate. Front Tarsal Claws: Symmetrical. Male Genitalia (Figures 343-345): As in P. ornaticollis except combined length of paramere and phallobase 1.6 times longer than phallobasic apodeme, paramere apex more acuminate, dorsal and ventral sinus narrower, and phallic plates not abruptly widened at middle. NATURAL HISTORY.—Unknown except the holotype was collected in December. ETYMOLOGY.—Latin noun sinus (pocket) + noun apex (tip) + adjectival ending -is, referring to the emargination of elytral apex.

LOCALITY RECORDS (Figure 369).—I examined 1 adult specimen from South America. VENEZUELA: Estado de Cara- bobo: San Esteban (FMNH, 1 male, holotype).

49. Perilypus fioralis (Gorham), new combination

FIGURES 333. 340-342, 346, 347, 368

Colyphus fioralis Gorham, 1882:142 [holotype: male, deposited in BMNH; type-locality: Volcan de Chiriquf, Provincia de Chiriqui, Panama]. DIAGNOSIS.—Distinguishable from all other congeneric species by the unique configuration of the elytral vittae. The flavous discal vitta begins at the elytral midbase and arches to join the epipleural margin vitta at the elytral apical fifth (Figure 342). DESCRIPTION.—Form: As in Figure 342. Size: Length: males, average about 6.8 mm, range 6.6-6.9 mm; females, average about 6.7 mm, range 6.0-7.4 mm. Width: males, 2.3 mm; females, average 355 about 2.1 mm, range 1.9—2.4 mm. Two males and 3 females measured. Color: Clypeus, frons, posterior epicranial region, FIGURE 355.—Perilypus galbeus, habitus. (Scale = 1 mm) venter of head, and antenna flavotestaceous; an- 114 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY terior epicranial region and gena black; prothorax bution extends from Provincia de Chiriqui, to flavotestaceous, except pronotum with black mid- Provincia de Panama, Panama. basal macula; legs flavotestaceous; meso- and LOCALITY RECORDS (Figure 368).—I examined 5 adult speci- metasternum black; elytron predominantly black, mens from Central America. PANAMA: Provincia de Chiri- with sinuous discal and linear marginal vitta con- qui: Volcan de Chiriqui (BMNH, 2 males, holotype; GEKI, 1 verging at apical fifth; abdomen black. female; MNHP, 1 female). Provincia de Panama: Cerro Head: Interocular depression, frontal umbo, and Campana (HFHO, 1 female). eye convexity as in P. ornaticollis; frons broad (HW/1OW, average about 2.0, range 1.8-2.1; 5 Immature Specimens specimens measured); antenna (Figure 333) distinctly serrate; length/width ratio of each male I studied two larvae and one pupa of P. limbatus antennal article 1.8:1.3:1.3:1.8:1.7:1.5:1.5:1.7:1.7: (Gorham), the only known immature specimens of 1.2:2.0; AL/PLS sex dimorphic, male 2.6, female Pcrilypus. Although the larval description is based 2.3. on one species, I tentatively consider the description Pronotum: Boldly transverse (PL/PW, average generic even though the specimens studied prob- about 0.78, range 0.71-0.80; 5 specimens measured); ably do not present adequately the full complement subapical depression deeply impressed; side margin of generic character states. Fortunately, the com- of pronotum proper boldly convex; fovea promi- prehensive publication of clerid larvae by Boving and Champlain (1920) provides some basis for nent, broad and shallow; prebasal depression shal- making tentative decisions about generic charac- low; pronotal collar narrow. terizations, even when only a few larval specimens Mesoscutellurn: Transverse. are available. Elytron: Elytral surface as in P. ornaticollis Lack of sufficient data and specimens precludes except surface smooth, shiny, less setose, and punc- a precise age determination of the two larvae tations reduced near suture and apex; epipleural examined. On the basis of size, the larger specimen fold very broad and convex (as wide as antennal probably belongs to at least the penultimate larval article 11); apex feebly sinuatotruncate. instar. Front Tar sal Claws: Symmetrical in both sexes. I do not provide a description of the pupa at Male Genitalia (Figures 340, 341): As in P. this time because the only available specimen exhib- ornaticollis except phallobasic apodeme as long as its most of the Perilypus imaginal characteristics. phallobase and paramere combined; dorsal and The small uncinate urogomphi, and chaetotaxy, ventral sinus broader, latter only slightly shorter may be taxonomically important, but I cannot than ventral; and phallus uniformly slender (2 decide without comparative information. In lieu specimens examined). of a written description, habitus figures of the Female Genitalia (Figures 346, 347): As in P. pupa (Figures 380, 381) are provided as prelimi- ornaticollis except ventral lobes more prominent nary diagnostic aids. and proctiger reduced (1 specimen examined). DIAGNOSIS.—Larval Perilypus are distinguished Internal Reproductive Organs: Male, anterior from larval specimens of most other genera of accessory gland uniramous, coiled throughout (1 Clerinae studied to date by having only one seta on specimen examined). the outer surface of the mandible. The only known VARIATION.—Antennal length is sex dimorphic, exceptions are Trichodes bicinctus Green and T. being longer in males than in females (AL/PL, nutalli (Kirby) (Foster, 1976); the conspicuously male 2.6, females 2.3). The elytral vitta varies in concave hypostomal margin in Trichodes distin- width. guish members of that genus from the available NATURAL HISTORY.—In August, at 915 meters on larvae of Perilypus in which the hypostomal margin Cerro Campana, H. and A. Howden collected 1 is feebly oblique. Foster (1976) provided detailed adult specimen by beating. G. C. Champion col- characterizations of Trichodes larvae in his revision lected 4 specimens from Volcan de Chiriqui at of North American Trichodes. The mandible of elevations from 915 and 1220 meters. other known clerine larvae has at least two mandib- DISTRIBUTION (Figure 368).—The known distri- ular setae. Also, the anterior margin of the labrum NUMBER 25J7 115 is emarginated deeply in larvae of Perilypus; the article 2 elongate and feebly arcuate. Labrum (Fig- anterior labial margin in other known clerine ure 375) transparent and well developed; side larvae is more transverse, and shallowly emargin- margins serrulated; anterior margin strongly emar- ated or feebly arcuate. ginated; aboral surface bearing eight large setae DESCRIPTION.—Form (Figure 370): Digitiform, arranged into two anterolateral groups of four. and cylindrical except head capsule flattened dorso- Epipharynx (Figure 375) with anterior row of 12 ventrally; forebody more tapered than hindbody; truncate epipharyngeal seta, and four epipharyngeal densely vested with pale setae. sensilla; latter dorsad to tormal sclerite, which Size: Length 8.8 mm, width 1.6 mm. extends into elongate processes posterolaterally. Color: Mandible, cranium, pronotal plate, and Thorax (Figures 370, 378): Prothorax with notal ninth abdominal segment dark brown; remainder plate strongly sclerotized, scutiform, and divided of mouthparts, antennae, legs, prosternal plate, and by a light midlongitudinal line; prosternum (Fig- meso- and metanotal plates light brown; abdominal ure 378) with elliptical midlongitudinal prosternal terga with pair of transverse purpurescent maculae; plate (the sternal plate of Boving and Champlain, remainder of body white. 1920:581) that adjoins two transverse lateral pro- Head (Figures 371, 372): Prognathus and prosternal plates (the presternal area of same authors). truding. Head capsule transverse (length 0.9 mm, Meso- and metathorax with pair of feebly scle- width 1.0 mm) plane, venter feebly tumescent; rotized notal sclerotizations; sternomedial scleroti- epicranial suture absent; frontal sutures feebly zation elliptical in mesothorax and subovate in arcuate at anterior third; frons triangular, with metathorax, plates feebly sclerotized in both seg- shallow longitudinal rugosities; epicranium smooth; ments. Legs well sclerotized, composed of five maxillary hypostomal margin (Figure 371) feebly segments that taper to claw; coxa rectangulate; oblique; gular sutures feebly sinuous; gular plate femur as long as longitudinal diameter of coxa; narrow, tapering posteriorly. Five stemmata (Figure tibia narrow, about as long as femur; tarsus 372) present; stemmata aligned in two parallel unguiform. vertical rows, three in anterior row and two in Abdomen (Figure 370): Segments 1 to 8 without posterior row. Antenna (Figure 376) composed of ampullae or sclerotizations; segment 9 (Figure 379) three articles and broad transparent basal mem- with strongly sclerotized notal plate, which projects brane; article 1 largest, about twice as long as wide; as pair of widely separated uncinate urogomphi; article 2 quadrate, ventroapical region projecting segment 10 reduced, entirely covered by segment 9. a conical antennal appendix; article 3 about three VARIATION.—In addition to size, the smaller times as long as wide and with long stout apical larva (length 8.0 mm, width 1.9 mm) differs from seta. Mandible (Figure 373) falciform, with one seta the larger one most conspicuously by the bluntness on posterior outer surface; mandibular groove of the mandibular apex (Figure 374), shallower deeply invaginated; apex unidentate; retinaculum mandibular groove, equilateral head capsule, and shallow, present on middle of masticatory surface; by having shorter integumental setae. lacinia mandibulae present, setiform, and only DISCUSSION.—In addition to the immature speci- feebly bifid apically. Maxilla (Figure 377) with mens of Perilypus described in this paper, I studied cardo and stipes trapezoidal, each with oblique larvae of the genera Placopterus, Enoclerus, Than- transverse plate at base, plate of former asetose, of asimus, Trichodes, Opilo, and Tillicera. Because latter bisetose near outer margin; palpiger feebly larvae of only a few genera of Clerinae are avail- sclerotized, bisetose at inner margin; palpus with able, any discussion of relationships based on larval three articles successively narrowing from basal character states is obviously tentative. Nevertheless, article, article 1 transverse, article 2 quadrate, such a discussion is presented to stimulate resyn- article 3 elongate; pedunculate seta well developed. theses as additional data become available. Labium (Figure 371) with rudimentary ligula; sub- The relative apotypy and plesiotypy of Clerinae mentum and mentum faintly sclerotized, former larval characters is beyond surmise at present, elongate, latter transverse; palpiger strongly scle- because few clerine larvae are available for study. rotized, with one stout seta on inner apical margin; However, the presence of the lacinia mandibulae in palpus with two articles, article 1 subquadrate, the larvae of Perilypus (Figure 373) and Placopterus 116 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY (Figure 382) suggest a potentially close relationship. unnecessary to rediscuss them or to restate their The lacinia mandibulae has heretofore not been logic as justification for their use herein. However, reported in Cleridae although Crowson (1964:277) I want to emphasize, an aspect of Hennig's noted its occurrence in other families of the super- approach apparently not fully appreciated; that is, family Cleroidea. Unfortunately, I have not been systematic studies in which Hennigian methods are able to substantiate the possible close relationship applied result in a more comprehensive presentation between Perilypus and Placopterus on the basis of of available data about the organisms under study. adult synapotypy, but earlier (p. 20) I alluded to The reason for this should become evident in the the superficial similarities between some of their following paragraphs. members. Should the lacinia mandibulae be judged Fundamental to Hennig's methods of phylo- a synapotypic characteristic, then it would be quite genetic analysis are the breakdown of phenotypic legitimate to infer sister group relationship between resemblance into symplesiomorphy, synapomorphy, the aforementioned two genera since it is not and convergence (for definitions see Hennig, 1965: unusual for taxa to lose indicators of "immediate" 102), and the establishment of monophyletic groups ancestry in the imago stage while retaining them only on the basis of shared apomorphic character in the immature stages. This is precisely why states; that is, a taxon is monophyletic when it Hennig (1966:32) stressed the importance of the includes all the known descendents of that taxon's holomorphological approach in phylogenetic ancestor; the descendents are recognized by the systematics. concept of synapomorphy. The second item of interest regarding the afore- Hennig's approach to phylogenetic analysis has mentioned larvae is that they provide some hints of an inherent prerequisite for in-depth investigations relationships at suprageneric level. The configura- of characters from as many aspects of an animal's tion of the hypostomal margin separates all the gestalt as is available for study; hence, the possi- larvae examined into two groups. In Perilypus, bility of discovering synapomorphies is maximized. Placopterus, and Enoclerus the margin is feebly How readily such similarities are detected is, of oblique (as in Figure 371), whereas in the remaining course, largely dependent on a group's evolutionary genera the hypostomal margin is distinctly concave history. Often it is dependent on a specialist's (Boving and Champlain, 1920, fig. 36). When the willingness to conduct comprehensive analysis of closely related genera of each group are brought more than just the readily observable characters. into alignment an interesting pattern emerges, one In-depth studies of phenotype (morphological or that may have considerable heuristic value. The otherwise) are essential to the holomorphological dichotomy, based on hypostomal margin configura- approach of character analysis forwarded by Hennig tion, essentially separates the more recently evolved (1966:180). Results from such an approach are genera (Perilypus, Placopterus, Enoclerus) from the aforementioned older generic groups (Trichodes, presented in phylogenetic systems in which rela- etc). The former group of genera are exclusively tionships are recognized by definable criteria New World whereas the latter group is predomi- (autapomorphies or synapomorphies), which theo- nantly Old World except for Priocera, a New retically best approximate evolutionary history. The World genus with apparent Old World affinities. concept of monophyly by autapomorphy only is one of the crucial differences between the theory of phylogenetic systematics advocated by Hennig Phylogeny and Zoogeography and that championed by Mayr (1969), or Sokal and Sneath (1963). Works not mentioned elsewhere in PHYLOGENETIC METHODS this paper that discuss some of Hennig's phylogenetic principles include Brundin (1966, 1968, The postulated phylogenetic relationships within 1972), Tuomikoski (1967), Darlington (1970), Schlee Perilypus are illustrated in Figures 383-386 and (1971), Kavanaugh (1972), and Mayr (1974). are based on Hennigian principles (Hennig, 1965, CHARACTER PHYLOGENY.—The phylogenetic status 1966). Hennig's tenets for reconstructing phylogeny of character states, whether apotypic (derived) or were copiously reviewed elsewhere and I think it plesiotypic (primitive), must be resolved before NUMBER 227 117

P. insect US (exact loca

FICURE 356.—Distribution map of the frontalis and criocerides groups. sister-group relationships can be established (the criteria require of the specialist a broad under- prefixes apo- and plesio- are combined with the standing of character states (particularly their dis- neutral suffix -typic (cf. Ball, 1975), not with tribution) among diverse groups relevant to the -morphic which has strictly morphological implica- taxa under investigation. tions). Criteria for inferring character phylogenies CRITERION OF FREQUENCY OF OCCURRENCE.—This were proposed and/or reviewed in detail by Hennig is the most conventional method of predicting a (1965, 1966), Maslin (1952), Schlee (1969), Kluge characteristic's phylogenetic status, particularly and Faris (1969), Marx and Rabb (1970, 1972), when two-state characters are involved. The objec- Crowson (1970), Munroe (1974), Ross (1974), and tive is to determine how extensively a character by Ball (1975). Of the criteria mentioned by these state is distributed within the taxon under study authors those most relevant to this study are briefly (e.g., genus) and among taxa of at least the next discussed below. The application of most of these highest category (e.g., subtribe). Needless to say, SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 118

P. relucens »P.viridipennis P.levis

L^^S pjP.chaletoides -^~Vij £ P. bicolor 'A 1o

359

FIGURES 357-359.—Distribution maps: 357-358, the viridipennis group; 359, the chaletoides group. the more inclusive the survey of taxa the more tion their extensive distribution is a result of credible becomes the assessment of character phy- inheritance, not due to independent evolution. logeny. Character states widespread among diverse Conversely, character states with limited distribu- groups are classified as plesiotypic on the assump- tions are interpreted apotypic because their re- NUMBER 227 119

O P.prolixipenis v£) P.testaceicornis 0 P.decoris X P.latilira ^ Rlimbatus ® Rbuga ® P.iris © Rapocopatus M P.columbicus O P.acus

FIGURES 360-362.—Distribution maps of the limbatiis group. 120 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

.carbonarius ^yP.caliculus P.distinctus P.latilobus P.calcaris P.sapientis P.cultratus P. reventazon P.exilis

P. immitis P. orthopleuridus P. nigriventris P. claudus P.bilineatus P.ordinatus P. cerroazul P.crassus

FIGURES 363-365.—Distribution maps of the reventazon group. NUMBER. TZ1 121

P.bicristatus P.quadrilineatus P.telepboroides P. coroniformis £) P.fluetus

P.ornaticollis

P. ventral is P.antarius P.floralis P.galbeus Rprolixicornis P.pilatus

FIGURES 366-369.—Distribution maps: 366, the quadrilineatus group; 367-369, the ornaticollis group.

stricted occurrence is an assumed manifestation of gence is more common than character convergence. recent evolution. However, limited distribution of CRITERION OF CORRELATION TO TRANSFORMATION characteristics may also indicate massive extinctions SERIES.—Morphoclines, or morphological transfor- in lineages once rich in taxa. Should the majority mation series, in which the order and direction of of taxa of such a lineage become extinct, the few change is known, provide a means of predicting survivors would exhibit sparsely distributed, character phylogeny of less understood morpho- relictual characteristics; these could be mistaken as clines, or of states of two-state character systems recently evolved on the basis of the criterion under whose phylogenetic value is unpredictable by the consideration. While relictual characteristics, among criterion of frequency of occurrence. Maslin (1952) extant taxa, are generally considered the exception, and Hennig (1966) discussed various ways by which their potential presence must be carefully con- the relative apotypic and plesiotypic extremes of sidered when the criterion of frequency of occur- transformation series can be inferred. In general, rence is used. The application of this criterion, the most practical way is to establish the most and of some that follow, requires the assumption plesiotypic extreme by the criterion of frequency that in evolutionary progression character diver- of occurrence and infer the opposite extreme to be SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 122

HYPOSTOMAL MARGIN

GULA

STEMMA 372

374

MANDIBULAR GROOVE

LACINIA MANDIBULAE 0.5 mm FICURES 370-374.—Perilypus limbatus larva: 370, habitus; 371, head, ventral view; 372, head, lateral view; 373, mandible, older larva (see p. 114); 374, mandible, younger larva. (Scales = 1 mm) TRUNCATE EPIPHARYNGEAL 376 SETA EPIPHARYNGEAL SENSILLUM

^ANTENNAL APPENDIX

PEDUNCULATE LATERAL PRO- SETA STERNAL PLATE MID-LONGITUDINAL PROSTERNAL STIPES PLATE

CARDO

FIGURES 375-379.—Perilypus limbatus larvae: 375, epipharynx; 376, antenna; 377, maxilla; 378, thorax, ventral view; abdominal segments vm-ix, lateral view; 379, abdominal segment ix. (Scale = 1 mm) 124 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

K.D.

FIGURES 380-382.—Perilypus limbatus pupae: 380, lateral view; 381, ventral view. Placopterus thoracicus: 382, mandible. (Scales = 1 mm) NUMBER 227 125 apotypic. The state, or states, between the apotypic interpretation, the basic tenet involves a search for and plesiotypic extremes are considered phylo- plesiotypic states (particularly of transformation genetic intermediates. In Hennig's phylogenetic series) of structurally rich characters that in aggre- system, intermediate (or transitional) character gate are considered the basic or ancestral plan. It states sometimes provide the only clue for inferring is assumed that the relatively apotypic states of sister-group relationships (Erwin 1974, fig. 161). extant descendents ultimately evolved from that CRITERION OF CHOROLOGICAL PROGRESSION.—Chor- plan. The basic plan (or aggregate of plesiotypic ology may be used as an index of character state characteristics) is usually established by the plesio- classification only after the spatial origin of a taxon typic extreme of transformation series or by the has been postulated. When this has been done criterion of frequency of occurrence. The taxo- structural character states may be correlated with nomic level at which the ancestral plan is estab- distributional character states. In general, and in lished is, of course, dependent on the taxonomic the absence of contrary evidence, one assumes that level of the group under study and on the cate- characteristics of relictual taxa are plesiotypic and gorical level at which such a plan can be credibly that those of taxa from derivative geographical established. regions are apotypic. Hennig (1966:95) provided It may be particularly difficult to recognize the the terms apochory and plesiochory to express, basic plan in recently evolved groups or in those respectively, the relative derived and primitive dis- where the higher classification is not well under- tributional states in chorological transformation stood, because in such cases recognition and series. interpretation of transformation series may be CRITERION OF CORRELATION WITH ADAPTIVE SIG- particularly difficult. Concordant transformation NIFICANCE.—As noted by Munroe (1974:76), "in- series involving complex organ systems are par- ferred adaptive significance is usually present, but ticularly important in this method of establishing hidden, in discussions of the distribution of charac- character phylogeny; mainly because it is the ter states in taxa." While it is often difficult to plesiotypic extreme of such series that lends credi- postulate, much less demonstrate, the adaptive bility to postulations of what the basic plan is. significance of a substantial number of character states, most groups of organisms evolved some character complexes whose adaptive function is ZOOGEOGRAPHIC METHODS fairly obvious. Character states judged apotypic on While hypotheses about speciation and phylogeny the basis of adaptive significance can be correlated have been organized into comprehensive theoretical with characteristics whose adaptive significance is models (of species by Mayr [1942], of phylogeny by cryptic, or whose phenotypic expression may not Hennig [1966]), those about historical zoogeography directly reflect adaptation. The latter kind of remain unconsolidated. The Darwinian concepts characteristic may represent manifestations of of center of origin and of dispersal of species genetic mechanisms not related to "immediate" remain the most practical tools for predicting adaptive needs. For example, their expression may ancestral distributions. Croizat, et. al. (1974), re- be due to genetic pleiotrophy or based on genetic cently criticized these concepts and provided alter- recombination generated by drift. native guidelines for historical biogeography at The substantial number of mimetic patterns biotal level. (obviously adaptive) present in Cleridae will make According to Croizat, et al. (1974:265), "the gen- the criterion of correlation with adaptive signifi- eral features of modern biotic distributions have cance particularly useful for inferring character been determined by subdivisions of ancestral biotas phylogeny. Munroe (1974:76) has commented in response to changing geography." I do not reject about the credibility of such inferences involving the concepts discussed by these authors; indeed the characters pertinent to mimicry and to warning idea that contemporary macrobiotic distributions coloration. resulted from fragmentations of ancestral biotas is CRITERION OF DEVIATION FROM A BASIC PLAN.— quite consistant with past geologic and climatic Schlee (1969) and Munroe (1974) discussed this events, such as continental drift and the less cata- criterion in considerable detail. According to my strophic events leading to continental refugia. On 126 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY the other hand, I cannot accept their total rejection RELATIONSHIPS AMONG THE SPECIES GROUPS of the concept of center of origin and its corollary, dispersal of species, primarily because faunal frag- Figure 383 illustrates the proposed phylogeny mentation predicates faunal expansion (expansion of the species groups and Table 1 lists the charac- accomplished by various possible mechanisms of ter states on which the reconstruction is based. The dispersal). It seems to me that a highly vagile species scheme is provisional, as all proposed phylogenies is capable of at least fortuitous dispersal during its must be, and is weakened by several convergences history, particularly in regions such as Central and reversals. Nevertheless, 1 feel that the analysis America where vicissitudinous climates have broadly reflects what might have transpired during undoubtedly affected dispersal potentials. The the early history of Perilypus; at the very least a question that remains, therefore, is this: Which of foundation of relationships is presented, one that the aforementioned approaches to historical zooge- is open to the challenge of subsequent discoveries ography is most useful? I submit that both have and resynthesis. merit, but generally at different biotic and/or The Placopterus-Perilypus stock probably dif- taxonomic levels. Irrespective of biotic or taxonomic ferentiated in northern Central America perhaps level, it is the nature of the available data that during mid-Tertiary time when northern Central usually determines the approach to historical America was discontinuous with a series of volcanic zoogeography. islands (now southern Central America) and iso- In general, zoogeographic relationships within lated by a wide sea gap from South America Perilypus are discussed in terms of faunal limits (Malfait and Dinkelman, 1972, fig. 3; Ball, 1975, and vicariance, and because the distributions of most of the species remain poorly documented the treatise is brief and fragmentary. Postulations about historical zoogeography are based on the concept of center of origin and I assume that vicariance reflects dispersal. Distributions of the South Amer- ican species are correlated with Quaternary refugia postulated by Haffer (1969), and Williams (1970), 4,5^,7,12,24 and Brown (1975). 25,26,27

Phylogenetics and Zoogeography of Perilypus 1,16 l2b,8,15b

INTERGENERIC RELATIONSHIPS

Earlier in this paper (p. 115), I suggested the possible sister group relationship between the genera Placopterus and Perilypus, based primarily on the presence of the lacinia mandibulae in the larvae. I cannot offer additional statements regarding intergeneric kinships based on potential synapotypic similarity. Otherwise, I can comment that there are at least three Central American genera (Colyphus, Clerosoma, and Blaxina) whose phylogenetic placement probably belongs close to Perilypus and Placopterus. Perhaps it will be neces- 19,18,31 sary to first study the larval stages and adult internal anatomy before these genera are placed FICURE 383.—Proposed phylogeny of the Perilypus species groups (numbers refer to apotypic character states listed in into credible phylogenetic perspective. Table 1). NUMBER 227 127

TABLE I.—Plesiotypic and apotypic character states used in Figure 383 (references indicate the criteria by which the character states are judged apotypic)

Character state Number Character Plesiotypic Apotypic 1 $ody depth shallow (Fig. 260) deep1 Head 2 antenna clubbed (Fig. 80) serrate2 (Fig. 96) 2a moderately serrate 3 (Fig- 8) 21) boldy serrate • (Fig. 284) 3 articles 7-8 without microsetae (Fig. with microsetae "• * 60) (Fig. 19) 4 interocular depression not crescemic crescent ic' Thorax 5 subapical depression strongly impressed feebly impressed * depth 6 subapical depression strongly sinuous (Fig. 110) feebly sinuous l (Fig. 342) shape 7 mesoscutellum subquadrate transverse l Elytra 8 form rectangulate (Fig. 2) ovatex (Fig. 300) 9 puncture shape not favous favous * 10 puncture depth deep shallow' 11 puncture distribution throughout disc (Fig. 2) absent from apical third or more' 12 sutuial cleft not right angle (Fig. 27) right angle * (Fig. 309) l 13 microsculpture absent present (Fig. 307) 14 epipleural fold not convex convex • (Fig. 163) 15 apical slope gradual (Fig. 30) acute» (Fig. 29) 15b very gradual '• * (Fig. 31) 16 disc color predominantly black or predominantly castaneous piceous or testaceous * 17 discal vitta absent present * (Fig. 2) Legs 18 front trasal claws symmetrical asymmetrical ••4 19 Pygidium entire (Fig. 39) emarginate * (Fig. 94) Male genitalia Tegmen 20 pigmentation strongly pigmented feebly pigmented *• * 21 phallobase width normal (Fig. 82) very broad ' (Fig. 72) 22 parameral lateral depres- present absent *• • sion 23 parameral mesodorsal not denticulated denticulated ••" (Fig. 282) margin 24 phallobasic pigmental absent present •• • (Fig. 344) bands 25 combined length of para- more than 2 less than 2 «• * mere and phallobasic apodeme 26 First connecting membrane serrulated scales absent serrulated scales present' Phallus 27 phallic plate shape normal (Fig. 46) uniformly slender *• * (Fig. 296) 28 marginal denticles absent present *• * (Fig. 46) Female genitalia 29 dorsal lamina trilobed (Fig. 71) bilobed •• • (Fig. 49) 128 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Continued

Character state Number Character Plesiotypic Apotypic 30 coxital plates absent (Fig. 198) present • (Fig. 48) Male internal reproductive organs 31 anterior accessory gland not coiled coiled1 (Fig. 51) 32 Distribution northern Central America southern Central Americaa 32a South America * 1 Frequency of occurrence within Perilypus. 2 Frequency of occurrence within Clerinae. 1 Correlation with transformation series. * Correlation with adaptive significance. * Deviation from a basic plan. * Chorological progression.

fig. 119). In that region (north of Nicaragua) occur mentioned genital characteristics. Antennal serra- the more primitive species of Perilypus, all the tion undoubtedly developed to maximize surface species of Placopterus, and most species of the other area for forebody sensory organs and thus is a genera closely related to Perilypus. The hypothesis highly adaptive character, one that is probably of a Central American origin of Perilypus requires genetically complex and less likely lost or evolved the assumption of three independent southern dis- in parallel than genital-phallobasic characteristics persals, and the penetration into South America by (the reverse would have to be assumed if P. levis at least one species during each southern movement. were to be placed in the frontalis group). The Ancestral Perilypus evolved autapotypic favous diverging sister lineage of the frontalis stock evolved elytral punctations, asymmetrical front tarsal claws, apotypic phallic marginal denticles and female and coiled male anterior accessory glands. The genital bilobed dorsal lamina. The marginal den- second character state was subsequently lost in ticles were secondarily lost in P. levis and in the several lineages, that is, assuming its genotype is progenitor of the ornaticollis group. relatively uncomplicated and easily reversed; The ancestor of the criocerides lineage also otherwise one would have to make the unlikely remained in northern Central America, retained assumption that asymmetry of the front tarsal claws an antennal club, and acquired a nearly impunctate evolved independently in five out of eight major elytron and apotypic pygidium emarginate. Some lineages. Ancestral Perilypus differentiated into two reduction of elytron punctation occurred inde- lineages, one giving rise to the frontalis stock and pendently in the viridipennis group, although in the other to the basic stock of the remaining that taxon the loss of deeply impressed punctations species. The stem species of the frontalis group involved only the elytral apical third. The comple- apparently remained in northern Central America, mentary stem species evolved apotypic serrate an- evolved an apotypic feebly pigmented tegmen, and tenna and microsetigerous antennal articles 7 and very broad phallobase, and lost the parameral 8, and differentiated into the progenitor of the lateral depression. The first and third of these limbatus-viridipennis stock and the one for the characteristics are also present in specimens of remainder of the species. P. levis (viridipennis group) that differ most con- An ancestor of the limbatus-viridipennis line spicuously from members of the frontalis group by having apotypic boldly serrate antennae. In this dispersed to southern Central America (exempli- scheme, I assume that genital characteristics of fying apochory; page 125) and subsequently pene- P. levis evolved independently from those of the trated South America. It evolved a convex epi- frontalis group, and that antennal serration is more pleural fold, a characteristic retained by all likely to reveal patristic kinships than the afore- descendants except P. relucens; in that species the epipleural fold became grooved longitudinally. A NUMBER 227 129 slightly different convex epipleural fold is also the evolution of these species were (1) transforma- present in both species of the frontalis group and tion of a moderately deep and rectangular body in one species of each of the quadrilineatus and form to one that is conspicuously shallow and oval, ornaticollis groups. Therefore, convexity of the and (2) progression of antennal serration from epipleural fold may not be a strong indicator of moderately serrate to boldy serrate. Perfection of a patristic affinities, but, in the absence of additional Batesian mimetic interaction (probably applicable evidence contrary to the proposed grouping, I ten- to most of the mimetic species under consideration) tatively consider the presence of that characteristic often requires that the incipient mimic species in the limbatus-viridipennis assemblage an expres- evolve toward a model species morphologically and sion of monophyly. The monophyly of the limbatus ecologically. Thus the imitator and the model group is based on the presence of autapotypic usually look alike and occur together in places female coxital plates; the plates are only feebly where both are maximally exposed to the developed in the more primitive Central American selection agent (predator). These requirements species. From the stock of the limbatus group invite the assumption that ancestral quadrilineatus- evolved the viridipennis group whose members ornaticollis negotiated a generalized ecological shift share apotypic impunctate elytral apical third (or from a relatively compact vegetation assemblage to less). Although I found only one autapotypic char- a more open one. Once the shift to a lampyroid- acteristic for each of the limbatus and viridipennis type habitat was initiated genetically pliant species groups, the two lineages probably have been sepa- of Perilypus converged towards the lampyroid rated for a long time as the considerable habitus models and concomitantly differentiated. In this differences between their extant members suggest. paper, under "Natural History" (p. 4), I provide The progenitor of the remaining species groups, some evidence that an ecological shift did indeed a vicariant with the ancestor of the limbatus- occur during the history of Perilypus. viridipennis lineage, acquired apotypic moderately The monophyletic status of the quadrilineatus serrate antenna, the second of two intermediate stock is based on apotypic parameral mesodorsal states of a transformation series that begins with a denticulated margin, a characteristic independently plesiotypic clubbed antenna and ends with an evolved in one species of the limbatus, viridipennis, apotypic boldly serrate antenna. The ancestor of and reventazon groups. From the correlative ances- this lineage also evolved elytral microsculpture tral species evolved the geographically widespread and subsequently differentiated, producing the ornaticollis group, the most derived species group reventazon lineage and the stock ancestral to the in the genus. The species of that group share the chaleloides, quadrilineatus, and ornaticollis groups. following apotypic characteristics not duplicated The evidence that the reventazon lineage is elsewhere in Perilypus: crescentic interocular de- monophyletic is that its members share the apotypic pression; feebly sinuous pronotal subapical depres- characteristics of an acute elytral-apical slope and sion; transverse mesoscutellum; right angled an elytral discal vitta. Exceptions involving char- elytral sutural cleft; presence of phallobasic pig- acter reversals involve P. orthopleuridus whose mental bands; combined length of paramere and elytral apical slope is gradual, and P. immitis and phallobase over length of phallobasic apodeme less P. crassus in which the elytral disc is apparently than 2; first connecting membrane with serrulated avittate (I studied only a few specimens of the last scales; and a uniformly slender phallic-plate shape. two species). The lineage complementary to ances- The extent of derivitiveness and the widespread tral reventazon evolved shallow elytral punctations distribution of the ornaticollis group gives the im- and subsequently differentiated into the basic stock pression that the group is much older than its of the chaletoides lineage and the quadrilineatus- phylogenetic placement suggests. But the derivitive- ornaticollis lineage. ness of the ornaticollis group may also be Ancestral chaletoides developed a deep body and interpreted as a manifestation of accelerated diver- predominantly castaneous elytral color. The pro- sification, particularly when considering the positive genitor of the complementary stock established two influence that ecological shifts might have on diverse lines of mimetic species. The most obvious evolutionary rate. The extensive geographic range anatomical reconstructions that materialized during of the group is probably a reflection of the general 130 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY mimetic interaction between ornaticollis species three of the larger species groups, namely the and a variety of lampyroid models. A generalized limbatus, quadrilineatus, and ornaticollis groups. mimic is not restricted to the range of only one The reventazon, frontalis, criocerides, viridipennis, model species as a more specialized mimic might and chaletoides groups are treated briefly and in be. Therefore, the generalist can overlap the geo- discussion form only. graphical range (become widespread) of several The two species of the frontalis group are very model species without losing the protective benefits similar anatomically, and probably are recently of the mimetic interaction. derived: the species are sympatric in Guatemala and in El Salvador, but P. frontalis extends northward RELATIONSHIPS WITHIN THE SPECIES GROUPS into southern Mexico and P. sensilis southward into western Nicaragua; the two species may be Patristic and geographic relationships within partial latitudinal complements. Perilypus frontalis each of the species groups are discussed and/or evolved an apotypic boldly convex female epipleu- illustrated to whatever extent the data allow ron and boldly asymmetrical male front tarsal claws. (Table 2). A fairly complete treatise is possible for Perilypus sensilis lost the asymmetry of the front

TABLE 2.—Plesiotypic and apotypic character states used in Figures 384-386 (references indicate the criteria by which the character states are judged apotypic)

Character state Number Character Plesiotypic Apotypic Head 33 gular process rectangulate filiform * (Fig. 308) 34 antennal article 11 not as long as articles 9 as long or longer than and 10 combined articles 9 and 10 combined' Pronotum 35 discal vitta not maculated maculatedl (Fig. 293) 35a not furcate furcate' (Fig. 322) 36 discal concavities absent presentJ (Fig. 300) 37 macrosculpture absent present* 38 transverse wrinkles absent present • 39 Mesoscutellum not stalked stalked • (Fig. 310) 39a subquadrate triangular * 40 Leg color femora concolorous or front and middle femur bicolorous concolorous, hind femur bicolorous *• * Elytra Disc 41 contour plane or feebly convex undulated *•4 (Fig. 293) 41a not bicristate bicristate * (Fig. 300) 42 fascia absent present * Margins 43 base or posthumeral avittate with short vitta • (Fig. 287) 44 apical arcuate truncate *• • 44a not sinuatotruncate sinuatotruncate '•' 44b not upcurved upcurved1 Epipleural fold 45 contour plane obliquely convex • 46 position lateral ventrolateral' 47 dorsal margin not explanate explanate * 48 narrow longitudinal not acutely deflexed acutely deflexed •• * region proximal to epipleural fold NUMBER 227

TABLE 2.—Continued

Character state Numbev Character Plesiotypic Apotypic 49 Vertical seta normal exceptionally stout' Male genitalia 50 phallic plate not abruptly broadened abruptly broadened medially medially! (Fig. 328) 51 phallic plate marginal far removed from phallic proximal to phallic denticles apex apex' (Fig. 130) 52 phallic apex normal acute 53 dorsal sinus normal reduced '•" (Fig. 338) 54 phallobase not explanate explanate1 (Fig. 134) Female genitalia Ventral lamina 55 length normal (Fig. 131) short'(Fig. 140) 56 size normal (Fig. 152) minute2 (Fig. 150) 57 number of lobes three one1 Dorsal lamina 58 median lobe absent presentl 59 number of lobes three seven' Bacculi 60 dorsal bacculus not contiguous (Fig. 151) contiguous1 (Fig. 171) 61 contiguous dorsal bacculus narrow expandedJ (Fig. 180) 62 oblique bacculus not fused with coxital fused with coxital plate plateJ (Fig. 140) 1 Frequency of occurrence within Perilypus. * Frequency of occurrence within Clerinae. 1 Correlation with transformation series. 4 Correlation with adaptive significance. 5 Deviation from a basic plan. " Chorological progression.

tarsal claws and acquired, in convergence, an apotypic convex epipleural fold. In the criocerides group, P. criocerides probably ranges into southeastern Mexico; no exact locality in Mexico is available for its sister species, P. insectus. Except for the autapotypic characteristics that define the group, the two species do not show a close relationship to each other. The substantial habitus differences between them probably reflects the mimetic interaction between P. criocerides and an unidentified species of leaf beetle. Perilypus criocerides evolved apotypic denticulatad outer margin of phallic plate. My views of the relationships within the limbatus group are illustrated in Figure 284. In the group, a monophyletic group of four Central American spe- 130 cies is vicarious with its South American sister group. The progenitor of the Central American FIGURE 384.—Proposed phytogeny of the limbatus species vicarient evolved apotypic continguous dorsal group (numbers refer to apotypic character states listed in baculi, a characteristic convergent in P. reventazon. Tables 1 and 2). 132 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Further differentiation of the lineage established of the reventazon group and in P. virdipennis of the basic stock of P. decoris characterized by an the viridipennis group. In specimens of these spe- apotypic unilobed ventral lamina (also found in cies, however, the fascia are very variable in shape P. reventazon), and the sister stock characterized and may be poorly defined. This lineage subse- by an apotypic acute phallic apex. The latter stock quently evolved from an ancestral stock, in which subsequently gave rise to the latilira-prolixipenis the phallic marginal denticles (of the male geni- stock and the testaceicornis stock, which evolved talia) became more proximal to the phallic apex, specializations relevant to leg color. Also in P. and a basic stock that later differentiated into the testaceicornis the fused dorsal bacculi of the female apocopatus-iris and limbatus-columbicus lineages. genitalia became expanded in the posterior half. I I did not find an autapotypic character state for did not find a synapotypic characteristic to substan- the latter lineage which subsequently differentiated tiate the sister group relationship between P. lati- into two species. One of these, P. columbicus, lira and P. prolixipenis, but their anatomical simi- acquired an apotypic explanate phallobase. The larity suggest close relationship; the two species apocopatus-iris lineage is inferred to be mono- have probably not been reproductively isolated for phyletic because the member species share the very long. following synapotypic characteristics of the female The South American species of the limbatus genitalia: an oblique bacculus fused with coxital group are known only from the Andes ranging from plate and short ventral lamina. The remaining two Venezuela to Peru; P. apocopatus is vicarious with taxa, P. buga and P. acus, are probably sister spe- P. iris, and P. buga with P. acus. Perilypus limbatus cies, but this needs corraboration by synapotypy. is particularly widely distributed extending from An apotypic characteristic of the female genitalia northern Venezuela to southern Peru. The known present in specimens of P. buga is a dorsal lamina distributions of these species roughly coincide with with median lobe, and in those of P. acus a lateral Quaternary Andean forest refugia postulated by lobe of minute ventral lamina. Haffer (1969) and Vanzolini and Williams (1970). The sister taxon of the limbatus group is the Assuming that the coincidental distributions viridipennis group; in the group, P. viridipennis between limbatus group species and refugia have and P. relucens are vicariants. The other known some historical significance, we might assume that species of this group, P. levis, ranges widely, occur- the species in question arose in the Pleistocene ring in southwestern Colombia, across southern epoch when Andean forest refugia were presum- Venezuela, and eastward into the Guiana Highlands ably established. The paucity of external differ- of Brazil. The progenitor of the group diverged, ences among the species, suggesting Recent origin, producing the levis and viridipennis-relucens line- lends support to that hypothesis. Ancient refugial ages. This differentiation probably took place early systems, probably also forestal, might also have in the history of the group; the extant taxa of the influenced the evolution of the Central American two lineages are very different structurally. The members of the limbatus group. The known geo- levis lineage lost the phallic marginal denticles but graphic occurrence of these species similarly coin- evolved in convergence the boldly serrate antenna, cides with montane forests, such as those found on feebly pigmented male genitalia, and the absence of the Panamanian Chiriquf range and Cerro the lateral depression of the paramere, as well as Campana, which contain more or less coherent the nonconvergent characteristic spatulate tegmen. biotas. It seems likely that Pleistocene climatic I found no unequivocal autapotypic charactristic changes (Vuilleumier, 1971) which profoundly for the viridipennis-relucens stock, except perhaps influenced South American forests also markedly the predominantly refescent color that is present in affected forest development in southern Central most specimens of the two extant species. America. In the reventazon group several species range The South American species of the limbatus widely in northern Central America: P. carbonarius group are more structurally homogeneous exter- and P. caliculus occur in Mexico from Sonora to nally than their Central American relatives. The Oaxaca and apparently are quite abundant in the basic stock evolved elytral fascia, an apotypic char- Transvolcanic Sierra; P. orthopleuridus is most acteristic independently evolved in P. carbonarius prevalent in southeastern Mexico and extends into NUMBER 227 133 southern Honduras. Most of the other species the southwest and P. bicolor extends northward appear to be localized in highlands of either central to Durango. The ancestors of the chaletoides group or southern Mexico, central Guatemala, central produced two structurally similar species that are Costa Rica, or northern Panama; P. crassus is the markedly different in elytral color. Perilypus chale- only species of the group indigenous to South toides acquired a dark streak along the suture of America. an otherwise castaneous elytron while P. bicolor In aggregate the species of the reventazon group evolved totally stramineous elytral coloration. are very similar externally; consequently, little can In the quadrilineatus group, P. quadrilineatus be stated about their patristic relationships. The ranges widely in eastern Mexico, but the species species differ substantially in characteristics of the also extends into southern Guatemala; P. tele- genitalia, however, but because aedeagal differences phoroides is vicarious with P. bicristatus. Figure 385 are so pronounced genital character states are gen- illustrates my views of the phylogenetics of the erally unsuitable for establishing sister groups; thus quadrilineatus group. Perilypus quardilineatus, no phylogeny is postulated. Despite these short- characterized by an apotypic dorsal margin of the comings some brief statements about relationships explanate epipleural fold, was apparently the first can be made. offshoot of the ancestral stock. The rest of the The robustness of the male pygidium may be species are derivatives of an ancestor that acquired used to roughly distinguish two clusters of species. a flatter, more oval body and one that evolved the In specimens of P. orthopleuridus, P. calcaris, P. apotypic characteristic, narrow longitudinal region exilis, and P. rerroazul the pygidium is quite ro- proximal to the acutely deflexed elytral posthumeral bust (Figure 229) while in the remainder of the margin. This ancestor subsequently differentiated species it ranges from small (Figure 245) to mod- into the bicristatus-telephoroides and coroniformis- erately large (Figure 39). I cannot decide to what fluctus lineages. The progenitor of the first lineage extent these groupings reflect phyletic relation- attained a coarsely macrosculptured pronotum and ships, primarily because the pygidium of some of subsequently diverged, producing P. telephoroides the species, such as P. reventazon, is difficult to and the highly derivative P. bicristatus. Apotypic group according to size. Also difficult to interpret character states present in P. bicristatus include a are the various degrees of asymmetry of the front bicristate elytral disc, triangular scutellum, and tarsal claws, despite meristic comparisons. The pronotal discal concavities. Perilypus telephoroides front tarsal claws are exceptionally asymmetrical in P. orthopleuridus, P. calcaris, P. exilis, and P. immitis. There are two particularly discordant species in the reventazon group, P. orthopleuridus and P. crassus. Specimens of the first species differ most conspicuously from members of the other species of the reventazon group by the following character state reversals: gradual apical slope of the elytron and the nonarenose elytral surface. The sister species of P. orthopleuridus is probably P. calcaris; specimens of these species share an apotypic mesoventral margin of the emarginate paramere, an acuminate phallus, inflexed outer margin of the phallic plate and a very long female genital coxite. P. crassus, the only South American species of the group, is notably different because its specimens 23 have the antenna exceptionally serrate and the elytron truncate. FIGURE 385.—Proposed phylogeny of the quadrilineatus spe- The species of the chaletoides group are known cies group (numbers refer to apotypic character states listed only from Mexico; P. chaletoides are mostly from in Table 2). 134 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY acquired the derivatives, the vittate base of the coast, and follows a northern track that spans the posthumeral margin, and the transversely wrinkled Mississippi and Ohio River drainage systems. Peri- pronotum. The elytral surface became undulated lypus ventralis extends from north of the Isthmus in the complementary progenitor which later di- of Tehuantepec to central Guatemala; P. floralis verged into P. fiuctus and P. coroniformis; the lat- is vicarious with the South American P. sinuapicis. ter acquired an apotypic maculated pronotal vitta. The hypothetical phylogenetic reconstruction of the Elytral undulations evolved secondarily in P. ven- ornaticollis group is illustrated in Figure 386. The tralis of the ornaticollis group. most specialized species in the group, and perhaps In the ornaticollis group two species range widely. in the genus, is P. ventralis, which anatomically Perilypus ornaticollis occurs in southeastern Mex- stands far apart from other members of the orna- ico, extends northward along the Mexican east ticollis group. The apotypic character states that define this early derivative are: a filiform gular process, stalked mesoscutellum, heptalobed dorsal lamina, plus various modifications of the elytra including a ventrolateral epipleural fold and the convergent characteristic undulated elytral surface. Some of these character states of P. ventralis are undoubtedly anatomical specializations relevant to the mimetic existence of that species, particularly those of the elytra. The complementary stock of the P. ventralis lineage acquired a reduction of the dorsal sinus of the male genitalia and subsequently diverged producing the ornaticollis and the progenitor of the remaining species. The ornaticollis lineage evolved at least two apotypic characteristics, furcate pronotal vitta and an abruptly medially broadened phallic plate. Its sister stem species evolved a truncated elytral apex and subsequently differentiated into the prolixicornis-galbeus stock and a line in which the elytral apex became sinuatotruncate. Article 11 of the antenna became prolonged in the prolixicornis-galbeus line, which then differentiated into P. prolixicornis and P. galbeus. The first offshoot of the remaining stock was P. pilatus, which is characterized by apotypic robust elytral vertical setae. The remaining three species are derivatives from a progenitor that acquired an obliquely convex epipleural fold and one that later differentiated into the antarius and floralis-sinuapicis lineages. 4,5,6,7,12,24, |25,26,2> ' Perilypus antarius acquired an apotypic upcurved FIGURE 386.—Proposed phylogeny of the ornaticollis species elytral apex. I did not find an autapotypic character group (numbers refer to apotypic character states listed in state that may have been present in the ancestor of Table 2). P. floralis and P. sinuapicis. Abstract in Spanish

Se redefine, reclasifica e ilustra el g^nero americano Perilypus. Se sumariza la historia natural del grupo. Las especies prosperan, por los menos, en tres tipos de macrohabitat, que son los relativos al roble, a lianas y a plantas herbaceas. Los escarabajos de cuerpo rectangular habitan en robles y lianas, mientras que los de cuerpo ovalado viven en hierbas. Se asume que la correlaci6n habitat- perfil del cuerpo se debe a interacciones mime'ticas. Los Perilypus mime'ticos se consideran generalistas que en cuanto a caracterfsticas de aspecto y habitos han evolucionado simulando mas de un modelo repugnante. Las especies de Perilypus se encuentran desde el nivel del mar hasta los 3400 metros de altitud, pero la mayoria de ejemplares conocidos se colectaron entre 1000 y 2000 metros. Observaciones de laboratorio prueban que los escarabajos de Perilypus son extremamente predatores, siendo factores importantes en la limitation de su agresividad el tamaflo y la rigidez de las presuntas victimas. Se discuten los criterios usados en el reconocimiento de las especies y de los grupos infra y supra-gene'ricos; se describen las t£cnicas seguidas para las disec- ciones, ilustraciones y mediciones; se dan analisis morfologicos de los principales organos externos e internos, y se ofrece una clave de las especies y de sus grupos basada en caracteres de adultos. Las fases de larva y ninfa del ge"nero se describen por vez primera. Se dan los caracteres para cada grupo de especies. Para cada especie se dan sinonimia, combination diagn6stica, descripci6n, discusi6n de la variation estructural y cromatica, se dan sinonimia, combinaci6n diagn6stica, description, discusi6n de la variaci6n estructural y cromatica, discusidn de su historia natural, distribution geografica, derivation etimol6gica, lot alidades, notas generales e ilustraciones. Mapas de distribuci6n con simbolos facilitan la visualizaci6n de las areas geograficas de especies y sus grupos. En total se reconocen 49 especies en Perilypus y 30 de estas se describen como nuevas. Se establecen 12 nuevas sinonimias. Se postula una filoge'nesis de Perilypus usando el m£todo Hennigiano de analisis filogenetico. Se expone la zoogeografia del g£nero en t&rninos de lfmites faunisticos y vicarismo. Se discuten sumariamente conceptos de zoogeograffa historica ("rutas faunisticas", "centro de origen y dispersi6n" y "refugios for- estales") y dos de ellos se utilizan para explicar la distribuci6n de los taxa de Perilypus existentes. Las consideraciones sobre las afinidades entre diversos groupos varia segun los datos asequibles. Se presenta un estudio bastante com- pleto de los grupos mayores de limbatus, quadrilineatus y ornaticollis, en cambio el tratamiento del grupo reventazon, el mayor del g^nero, es muy fragmentario. Es probable que el Perilypus ancestral se desarroll6 en el norte de America Central durante el Terciario medio evoluci6nado al fin en ocho grupos espe- cificos. Los progenitores de los grupos frontalis, criocerides, chaletoides y quadrilineatus permanecieron en norte de Centro America asi como tambie"n sus descendientes. Los ancestros de los grupos limbatus y viridipennis se dispersaron hacia el sur de America Central; algunos miembros de ambos grupos penetraron ultimamente en Sur America. El grupo ornaticollis, el mas evoluci6nado en el ge"nero, y el grupo reventazon, especialmente, son de gran dispersi6n geografica.

135 Literature Cited

Ball, G. E. tera. Transactions of the Royal Entomological So- 1975. Pericaline Lebiini: Notes on Classification, a Syn- ciety of London, 94:273-310. opsis of the New World genera, and a Revision of 1964. A Review of the Classification of Cleroidea (Coleop- the Genus Phloeoxena Chaudoir (Coleoptera: Cara- tera), with Descriptions of Two New Genera of bidae). Questiones Enlomologicae, 11 (2): 143-242. Peltidae and of Several New Larval Types. Trans- Boving, A. G., and A. B. Champlain actions of the Royal Entomological Society of Lon- 1920. Larval of North American Beetles of the Family don, 116(12):275-327. Cleridae. Proceedings of the United States National 1970. Classification and Biology. 350 pages. London: Heine- Museum, 57(2323):575-649. mann Educational Books. Brown, K. S., Jr. Darlington, P. J., Jr. 1975. Geographical Patterns of Evolution in Neotropical 1970. A Practical Criticism of Hennig-Brundin "Phylo- Lepidoptera: Systematics and Derivation of Known genetic Systematics" and Antarctic Biogeography. and New Heliconiini (Nymphalidae: Nymphaline). Systematic Zoology, 19(1):1—18. Journal of Entomology, series B, 44(3):201-242. Desmarest, E. Brundin, L. [I860.] Cleriens. Pages 226-279 of part 2 in J. C. Chenu, 1966. Transarctic Relationships and Their Significance as Encyclopedic d'Histoire Naturelle. Paris: Libraire Evidenced by Chironomid Midges: With a Mono- de Firmin-Didot et C". graph of the Subfamilies Podonominae and Aphro- Doyen, J. T. teniinae and the Austral Heptagyiae. Kongliga 1966. The Skeletal Anatomy of Tenebrio molitor (Coleop- Svenska Vetenskapsakadaniens Handlingar, (4)11:1- tera: Tenebrionidae). Miscellaneous Publications of 472. the Entomological Society of America, 5(3):102-150. 1968. Application of Phylogenetic Principles in Systematics Dury, C. and Evolutionary Theory. Pages 473—495 in T. 1906. Ecological Notes on Some Coleoptera of the Cin- 0rvig, editor, Proceedings of the 4th Nobel Sympo- cinnati Region including Seven New Species. Jour- sium: Current Problems of Lower Vertebrate Phyto- nal of the Cincinnati Society of Natural History, geny. Stockholm: Almqvist & Wiksell. 20:151-156. 1972. Evolution, Causal Biology, and Classification. Zoo- Ehrlich, P. R., R. R. White, M. C. Singer, S. W. McKechnie, logica Scripta, 1:107-120. and L. E. Gilbert Campau, E. J. 1975. Checkerspot Butterflies: A Historical Perspective. 1940. The Morphology of Chauliognathus pennsylvanicus Science, 188(4185):221-228. (DeGeer) (Coleoptera: Cantharidae). Microentomol- Ekis, G. ogy, 5(3):57-85. 1975. Taxonomic and Nomenclatural Status of Clerid Chevrolat, M. A. Taxa Described by Massimiliano Spinola (1780- 1843. Coleopteres du Mexique. Magasin Zoologie, series 2 1857) (Coleoptera, Cleridae). Bollettino del Museo (Insectes), 5:1-26. di Zoologia dell'Universita di Torino, 1:1—80. 1874. Catalogue des Clerides de la collection de M. Chev- Ekis, G., and A. P. Gupta rolat. Revue et Magasin de Zoologie, series 3, 1971. Digestive System of Cleridae (Coleoptera). Interna- 2(6):252-329. tional Journal of Insect Morphology and Em- 1876. Mimoire stir la famille des Cle'rites. 51 pages. Paris. bryology, l(l):51-86. Cholodkovsky, N. Erwin, T. L. 1913. Necrobia ruficollis in St. Petersburg. Zoologischer 1974. Studies of the Subtribe Tachyna (Coleoptera: Cara- Ameiger, 42(12):529-531. bidae: Bembidiini), Part II: A Revision of the New Corporaal, J. B. World-Australian Genus Pericompsus Le Conte. 1948. Nineteen Notes on Systematics and Synonymy. Ento- Smithsonian Contributions to Zoology, 162:1-96. mologische Berichten, 12(281 ):243-246. Evans, M. E. G. Croizat. L.. G. Nelson, and D. Rosen 1961. The Musculature and Reproductive Systems of 1974. Center of Origin and Related Concepts. Systematic Atomaria ruficornis (Marsham) (Coleoptera, Crypto- Zoology, 23(2): 265-287. phagidae). Transactions of the Royal Society of Edin- Crowson, R. A. burg, 64(14):297-399. 1944. Further Studies on the Metendosternite in Coleop- Ferris, G. F.

136 NUMBER 227 137

1940. The Morphology of Plega signata (Hagen) (Neurop- 1969. The Ground-Beetles (Carabidae, excl. Cicindelinae) tera: Mantispidae). Microentomotogy, 5(2):33-49. of Canada and Alaska, Part I. Opuscula Ento- Forbes, W. T. M. mologica (Lund), Supplement 35:I-XIVII. 1925. The Second Abdominal Pleurite in the Higher Malfait, B. T., and M. G. Dinkelman Coleoptera. Psyche, 32(6):290-292. 1972. Circum-Caribbean Tectonic and Igneous Activity, Foster, D. E. and the Evolution of the Caribbean Plate. Bulletin 1976. Revision of North American Trichodes (Coleop- of the Geological Society of America, 83:251-272. tera: Cleridae). Special Publications of the Museum, Marx, H., and G. B. Rabb Texas Tech University, 11:1-86. 1970. Character Analysis: An Empirical Approach Applied Gemminger, M., and E. Harold to Advanced Snakes. Journal of Zoology (London), 1869. Familia XL1II, Cleridae. Pages 1722-1759 of volume 161:525-548. 6 in Gemminger and Harold, Catalogus Coleoptero- 1972. Phyletic Analysis of Fifty Characters of Advanced rum. Monaco: Sumptu E. H. Gummi. Snakes. Fieldiana Zoology, 63:1-11. Gorham, H. S. Maslin, T. P. 1878. Descriptions of New Genera and Species of Cleridae, 1952. Morphological Criteria of Phylogenetic Relation- with Notes on the Genera and Corrections of ships. Systematic Zoology, l(2):49-70. Synonymy. Transactions of the Entomological So- Matsuda, R. ciety of London, 1878(2):153-167. 1970. Morphology and Evolution of the Insect Thorax. 1882. Fam. Cleridae. Pages 129-168 in part 2 (Coleoptera) Memoirs of the Entomological Society of Canada, of volume 3 in Godman and Salvin, editors, Biologia 76:1-431. Centrali-A mericana. Mayr, E. 1886. Supplement: Colyphus-Poecilochroa. Pages 335-337 1942. Systematics and the Origin of Species. 334 pages. in part 2 (Coleoptera) of volume 3 in Godman and New York: Columbia University Press. Salvin, editors, Biologia Centrali-Americana. 1969. Principles of Systematic Zoology. 328 pages. New Haffei, J. York: McGraw-Hill. 1969. Speciation in Amazonian Forest Birds. Science, 1974. Cladistic Analysis or Cladistic Classification? Zeit- 165:131-137. schrift fur zoologische Systematik und Evolutions- Hennig, W. jorschung, 12:94-128. 1965. Phylogenetic Systematics. Annual Review of Ento- Munroe, D. D. mology, 10:97-116. 1974. The Systematics, Phylogeny, and Zoogeography of 1966. Phylogenetic Systematics. 263 pages. Urbana: Uni- Symmerus Walker and Australosymmerus Freeman versity of Illinois Press. (Diptera: Mycetophilidae: Ditomyiinae). Memoirs of Hlavac, T. F. the Entomological Society of Canada, 93:1-183. 1972. The Prothorax of Coleoptera: Origin, Major Fea- Pic, M. tures of Variation. Psyche, 79(3): 123-149. 1941. Opuscula martialis, II. "L'Echange," Revue Lin- Holdridge, L. R., W. C. Grenke, W. H. Hathevvay, T. Liang, neene (16 April). [See Cleridae listing on pages 9-13.] and J. A. Tosi, Jr. Ross, H. H. 1971. Forest Environments in Tropical Life Zones: A 1974. Biological Syste/natics. 345 pages. Reading, Mass.: Pilot Study. 747 pages. New York and Toronto: Addison-Wesley Publication Co. Inc. Pergamon Press, Oxford. Schaeffer, C. Kavanaugh, D. H. 1904. New Coleoptera. Journal of the New York Ento- 1972. Hennig's Principles and Methods of Phylogenetic mological Society, 12:215-224. Systematics. The Biologist, 54(3): 115-127. Schenkling, S. Kluge, A. G., and J. S. Faris 1898. Zehn neue Cleriden nebst Bemerkungen iiber schon 1969. Quantitative Phyletics and the Evolution of An- beschriebene Arten. Deutsche Entomologische Zeit- urans. Systematic Zoology, 18(1): 1-32. schrift, 1898(2):361-368. Lacordaire, J. T. 1903. Coleoptera Malacodermata: Fam. Cleridae. Fascicle 1857. Famille XLI: Clerides. Pages 415-496 of volume.4 in 13 in P. Wytsman, Genera Insectorum. Bruxelles. Lacordaire, Histoire Naturelle des Insectes: Genera 1906. Die Cleriden des Deutschen Entomologischen des Coleopteres. Paris: Librairie Encyclopedique de National-Museums, nebst Beschreibungen neuer Roret. Arten. Deutsche Entomologische Zeitschrift, 1:241— LeConte, J. L. 320. 1880. Short Studies of North American Coleoptera. Trans- Schlee, D. actions of the Entomological Society of America, 8:194-195. 1969. Hennig's Principle of Phylogenetic Systematics, an Lindroth, C. H. "Intuitive statistico-phenetic Taxonomy"? Systematic 1957. The Principle Terms Used for Male and Female Zoology, 18(1): 127-134. Genitalia in Coleoptera. Opuscula Entomologica, 1971. Die Rekonstruktion der Phylogenese mil Hennig's 22:241-256. Prinzip. 62 pages. Frankfurt. 138 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Scotl, H. Bollettino del R. Laboratorio di entomologia agraria 1919. Notes on the Biology of Necrobia ruficollis, Fabr. di Portici, 16:49-140. (Coleoptera, Cleridae.) The Annals of Applied Tuomikoski, R. Biology, 6(2-3): 101-115. 1967. Notes on Some Principles of Phylogenetic Sys- Snodgrass, R. E. tematics. Annals Entomologici Fennici, 33(3):137- 1935. Principles of Insect Morphology. 667 pages. New 147. York: McGraw Hill. Vanzolini, P. E., and E. E. Williams Sokal, R., and P. Sneath 1970. South American Anoles: The Geographic Differ- 1963. Principles of Numerical Taxonomy. 359 pages. San entiation and Evolution of the Anolis chrysolepis Francisco: Freeman, 359 pp. Species Group (Sauria, Iguanidae). Arquivos de Solervicens, A. J. Zoologia Sao Paulo, 19:1-298. 1973a. El genero Epiclines en Chile. Annales Museo His- Vuilleumier, B. S. toria Natural Valparaiso, 6:161-186. 1971. Pleistocene Changes in the Fauna and Flora of South America. Science, 173:771-780. 1973. A Revision del genero Natalis Castelnau (Coleop- Whitehead, D. R. tera-Cleridae-Clerinae). Revista Chilena Entomo- 1972. Classification, Phylogeny, and Zoogeography of logia, 7:233-247. Schizogenius Putzeys (Coleoptera: Carabidae: Scar- Spinola, M. atini). Quaestiones Entomologicae, 8:131-348. 1841. Monographie des Terediles. Revue Zoologique par Wolcott, A. B. La Socie'te Cuxnerienne, 4:70-76. 1910a. Family XLII, Cleridae: The Checkered Beetles. 1844a. Essai Monographique sur les Cle'rites: Insectes In W. S. Blatchley, An Illustrated Descriptive Cata- Coleopteres. Volume 1, 386 pages. Genes. logue of the Coleoptera or Beetles (Exclusive of the 1844b. Essai Monographique sur les Cle'rites: Insectes Rhynchophora) Known To Occur in Indiana. Bul- Coleopteres. Volume 2. 216 pages. Genes. letin of the Indiana Department of Geology and Tanner, V. M. Natural Resources, 1:846-862. 1927. A Preliminary Study of the Genitalia of the Female 1910b. Notes on Some Cleridae of Middle and North Coleoptera. Transactions of the American Entomo- America with Descriptions of New Species. Zoologi- logical Society, 53:5-50. cal Series (Field Museum of Natural History Pub- Thomson, M. S. lication 144), 7(10):339-401. 1860. Materiaux pour servira une monographie nouvelle 1927a. A Review of the Cleridae of Costa Rica. Coleoptero- de la famille des Cle'rides. Musee Scientifique, logical Contributions, l(l):l-104. 1860:46-67. 1927b. Descriptions of a New Genus and Four New Species Tremblay, E. of American Cleridae. Coleopterological Contribu- 1958. Studio morpho-biologico sulla Necrobia rufipes DeG. tions, 1(1): 105-110.

REQUIREMENTS FOR SMITHSONIAN SERIES PUBLICATION Manuscripts intended for series publication receive substantive review within their originating Smithsonian museums or offices and are submitted to the Smithsonian Institution Press with approval of the appropriate museum authority on Form SI-36. Requests for special treatment—use of color, foldouts, casebound covers, etc.—require, on the same form, the added approval of designated committees or museum directors. Review of manuscripts and art by the Press for requirements of series format and style, completeness and clarity of copy, and arrangement of all material, as outlined below, will govern, within the judgment of the Press, acceptance or rejection of the manuscripts and art. Copy must be typewritten, double-spaced, on one side of standard white bond paper, with lVi" margins, submitted as ribbon copy (not carbon or xerox), in loose sheets (not stapled or bound), and accompanied by original art. Minimum acceptable length is 30 pages. Front matter (preceding the text) should include: title page with only title and author and no other information, abstract page with author/title/series/etc., following the established format, table of contents with indents reflecting the heads and structure of the paper. First page of text should carry the title and author at the top of the page and an unnum- bered footnote at the bottom consisting of author's name and professional mailing address. Center heads of whatever level should be typed with initial caps of major words, with extra space above and below the head, but with no other preparation (such as all caps or underline). Run-in paragraph heads should use period/dashes or colons as necessary. Tabulations within text (lists of data, often in parallel columns) can be typed on the text page where they occur, but they should not contain rules or formal, numbered table heads. Formal tables (numbered, with table heads, boxheads, stubs, rules) should be submitted as camera copy, but the author must contact the series section of the Press for edito- rial attention and preparation assistance before final typing of this matter. Taxonomic keys in natural history papers should use the alined-couplet form in the zoology and paleobiology series and the multi-level indent form in the botany series. If cross-referencing is required between key and text, do not include page references within the key, but number the keyed-out taxa with their corresponding heads in the text. Synonymy in the zoology and paleobiology series must use the short form (taxon, author, yearpage), with a full reference at the end of the paper under "Literature Cited." For the botany series, the long form (taxon, author, abbreviated journal or book title, volume, page, year, with no reference in the "Literature Cited") is optional. Footnotes, when few in number, whether annotative or bibliographic, should be typed at the bottom of the text page on which the reference occurs. Extensive notes must appear at the end of the text in a notes section. If bibliographic footnotes are required, use the short form (author/brief title/page) with the full reference in the bibliography. Text-reference system (author/year/page within the text, with the full reference in a "Literature Cited" at the end of the text) must be used in place of bibliographic footnotes in all scientific series and is strongly recommended in the history and technology series: "(Jones, 1910:122)" or ". . . Jones (1910:122)." Bibliography, depending upon use, is termed "References," "Selected References," or "Literature Cited." Spell out book, journal, and article titles, using initial caps in all major words. For capitalization of titles in foreign languages, follow the national practice of each language. Underline (for italics) book and journal titles. Use the colon-parentheses system for volume/number/page citations: "10(2):5-9." For alinement and arrangement of elements, follow the format of the series for which the manuscript is intended. Legends for illustrations must not be attached to the art nor included within the text but must be submitted at the end of the manuscript—with as many legends typed, double- spaced, to a page as convenient. Illustrations must not be included within the manuscript but must be submitted separately as original art (not copies). All illustrations (photographs, line drawings, maps, etc.) can be intermixed throughout the printed text. They should be termed Figures and should be numbered consecutively. If several "figures" are treated as components of a single larger figure, they should be designated by lowercase italic letters (underlined in copy) on the illustration, in the legend, and in text references: "Figure 9b." If illustrations are intended to be printed separately on coated stock following the text, they should be termed Plates and any components should be lettered as in figures: "Plate 9b." Keys to any symbols within an illustration should appear on the art and not in the legend. A few points of style: (1) Do not use periods after such abbreviations as "mm, ft, yds, USNM, NNE, AM, BC." (2) Use hyphens in spelled-out fractions: "two-thirds." (3) Spell out numbers "one" through "nine" in expository text, but use numerals in all other cases if possible. (4) Use the metric system of measurement, where possible, instead of the English system. (5) Use the decimal system, where possible, in place of fractions. (6) Use day/month/year sequence for dates: "9 April 1976." (7) For months in tabular list- ings or data sections, use three-letter abbreviations with no periods: "Jan, Mar, Jun," etc. Arrange and paginate sequentially EVERY sheet of manuscript—including ALL front matter and ALL legends, etc., at the back of the text—in the following order: (1) title page, (2) abstract, (3) table of contents, (4) foreword and/or preface, (5) text, (6) appendixes, (7) notes, (8) glossary, (9) bibliography, (10) index, (11) legends. i